SUMMARY
Intensive study of anting behavior in a captive Orchard Oriole (free of obvious ectoparasites) with worker ants of several species during a 31-month period showed the bird's basic anting pattern was similar to that described for most small species, but new or different in several aspects. To summarize present knowledge and for purpose of comparison, an analysis was made of all available anting records, including some unpublished materials. The compiled list of 148 species of anting birds includes 65 New World forms. Types of anting, theories concerning its significance, and the possible
relationship of anting to dust-bathing and sun-bathing are reviewed. Two basic types of this behavior are recognized: active anting (anointing by use of the bill) and passive anting (anointing by allowing ants to invade the plumage). Offered seven non-spraying ant forms, the Orchard Oriole exhibited consistent selectivity in choice of species for anting. Differences in odor, flavor and defensive mechanisms of acceptable and unacceptable ant species are described, the most important being that all acceptable ants (three species) were found to produce a burning sensation on the human tongue, whereas the four unacceptable ant forms did not. This effect cannot be ascribed to the non-caustic butyric acid believed to be produced by all ant species the oriole accepted. The thermogenic agent in these ants has not been determined. It appears that all ant species used by birds for anting may have thermogenic properties. Most species used are producers of formic acid, and may be presumed to be similarly stimulating. Since many substitute materials used for anting are also of this nature, "heat" seems to be the common factor in most, if not all, anting situations. A synopsis of the known and identified ants used by birds for anting indicates that birds in general are selective. With one exception (probably a conditioned response), birds seem to have used only species that either spray or exude repugnatorial fluids. The ability of spraying ants, at least Formica rufa and its allies, to eject a fine mist to a distance of 20 to 50 cm. is an important factor in anting that may not have been generally recognized. The Orchard Oriole applied ants by dabbing, not stroking. At the same time it rapidly vibrated its head. Regularly the bird anointed ventral surfaces of wing tips and of the base of the tail, and also the undertail coverts. Sides and flanks were touched only indirectly as the bird reached its vibrating head around toward wing or tail. Occasionally feathers of the belly and tibiae were anointed. Undertail coverts and bases of the rectrices received the most attention. Following the performance of anting with non-spraying ants, the bird was found to have ant odor on all these areas but not on other plumage tracts. The bird apparently crushed the ants before applying them. Crushing was found to increase the odor and thermogenic effect to the human tongue. Tripping and tumbling occurred regularly, resulting from interference of the tail, which was brought forward beneath the body during anting. Often the bird performed while clinging to the cage wall, at times anting in an up-side-down position. Sometimes the oriole gathered and held in the bill a large wad of ants before applying, discarding or eating it. Occasionally it selected and ate certain bits of ant-nest earth, possibly soil impregnated with ant exudations. Ordinary earth was not eaten.
Positive anting response was high in this bird. Of 80 contact days, it performed on at least 67. Negative response due to surfeit was rare. The longest period of successive daily performance was 10 days. Anting sessions, often lasting 45 minutes, became shorter as ants were offered daily. Physical stress of high intensity anting caused noticeable fatigue. Drowned, frozen, or heat-killed ants elicited pronounced anting reaction, except when the treatment caused loss of thermogenic property (i.e., through heat-kill or long-term freezing). The oriole distinguished variations among living worker ants of the colony, applying some individuals repeatedly, others only once, and discarding some without use. Taste-testing showed marked thermogenic variation among worker ants of a colony. Apparently the bird used most assiduously those individuals having strong heating value. This property of the ant seemed less pronounced in winter than in summer, and may be a cause of the bird's somewhat milder anting response in winter. The bird was not interested in ants primarily as food. Preening and bathing following anting were erratic and seemed caused as much by physical exertion and displaced plumage as by the possible effect of ant-ointment on the feathers. These non-spraying ants caused no unusual blinking or spreading of nictitating membrane, frequently reported in birds using spraying ants. Records of anting in birds obviously infested with ectoparasites appear to be extremely rare. Endoparasites as a factor in anting should be investigated. No report of an autopsy of an anting bird could be found, or even of micro-scopic examination for smaller ectoparasites. Literature examined failed to indicate that captivity either causes or suppresses anting or modifies significantly the specific anting patterns. The cause of anting in the Orchard Oriole was not determined. The bird's preferential treatment of the undertail region, and possibly the vent itself at times, was indicated by direct observation, by extreme displacement of undertail coverts, and by the fact that ant odor was strongest here. Close study showed this region received additional treatment when wing tips were anointed, due to juxtaposition of the wing tip with the tail. Although there was no evidence of gross ectoparasitism in the oriole, it is possible that ant substances relieved itching caused by minute ectoparasites or cysts of endoparasites, certain species of which are known to occur in skin of the undertail region of birds. Also, the bird appeared to derive sensual pleasure, possibly including sexual stimulation, from the thermogenic effect of the ants. The concept of sexual stimulation might throw light on several peculiar features sometimes seen in anting behavior, such as pressing and rubbing of vent upon the anting ground, drooping-wings posture following anting, convulsive shuddering and quivering in conjunction
with a double-wing-spread stance, quivering or shaking of wings and tail, individual differences in the amount and in the occurrence of anting among captives of a species, and the apparent irregularity with which birds seem to ant in nature. Several facts, derived from study of the literature and considerable reading in the field of myrmecology, may bear upon the problem. First, it is physically possible for ant substances to touch a bird's ventral posterior skin (including the vent) during either active or passive anting, due to ability of a bird to reach this area with its bill (shown in many anting descriptions and commonly seen in preening), and to the spraying force of ants. Secondly, passive anting with non-spraying ant species appears to be almost unknown; and, unlike active anting, it often entails raising of the contour plumage. Thirdly, while anointment of plumage and skin must become general when a bird permits spraying ants to invade its feathers in numbers, species (excepting grackles) that practice active anting apparently seldom or never apply ants to the anterior part of the body, the dorsal body or even to dorsal surfaces of quill feathers. Certain apparently aberrant anting motions, seen in the Orchard Oriole or reported in literature (incipient or desultory applications, anointment of unusual feather tracts), may possibly be due to inadequate amount of stimulant in the individual ant or in the substitute material used. The oriole's reactions to frozen and to heat-killed ants (stimulant reduced or destroyed) suggest correlation between the amount or degree of the thermogenic agent and the plumage area treated. This topic needs further investigation. In recognizing "heat" as the common anting factor, smoke must be given consideration as an anting substance. Smoke-bathing appears virtually unknown in the Western Hemisphere. Instances of species, even individuals, performing alike with ants, smoke, and burning or heated materials, all seem cases in point. This study demonstrates the importance of entomology to any solution of the anting problem. In common with others, it contributes little toward understanding why certain species of birds will ant while others apparently do not, but it does show that new species are continually being added to the list of anting birds. The fact that a species has not been reported does not necessarily mean that it does not ant. Likewise, it is risky to conclude that the anting response is absent in a species merely because individuals have refused to ant. Many questons raised here cannot be answered short of controlled experiments on a large scale. Perhaps this resume will provide an impetus for thoroughgoing experimentation, particularly with Common Crows (passive anting) and Blue Jays or Common Starlings (active anting), inasmuch as anting incidence seems to be unusually high among these species.
NCE Audub,on (1831:7) wrote of Wild Turkeys (Meleagris gallopavo)
rolling in 'deserted" ants' nests (Allen, 1946), and Gosse (1847:225)
reported Tinkling Grackles (Quiscalus niger) in nature anointing themselves
with lime fruits (Chisholm, 1944), an extensive literature on the anting ac-
tivities of birds has slowly evolved. The complete bibliography of anting
probably would approximate 250 items, yet the purpose of the behavior re-
mains unexplained.
Anting may be defined as the application of foreign substances to the plum-
age and possibly to the skin. These substances may be applied with the bill,
or the bird may "bathe" or posture among thronging ants which invest its
plumage.
Among numerous explanations for the use of ants are these: (1) the bird
wipes off ant acid, preparatory to eating the ant; (2) ants prey upon, and
their acids repel, ectoparasites; (3) ant acids have tonic or medicinal effects
on the skin of birds; (4) odor of ants attracts birds, much as dogs are drawn
to ordure or cats to catnip; (5) ants intoxicate the bird or give it unique
pleasurable effects; (6) ant substances on the plumage, irradiated by sun-
light, produce vitamin D, which the bird ingests during preening; (7) the
bird enjoys the movement of insects in its plumage; (8) ant substances pre-
vent over-drying of feather oils or give a proper surface film condition to
the feathers. For discussions of these possibilities, see Chisholm (1944, 1948:
163-175), Adlersparre (1936), IJzendoorn (1952a), Eichler (1936a), Klein-
schmidt (in Stresemann, 1935b), Lane (1951:163-177), Kelso (1946, 1949,
1950a, 1950b, 1955:37-39), Brackbill (1948), G6roudet (1948), Groskin
(1950), and McAtee (1938).
At least 24 kinds of ants and more than 40 substitute materials have been
used by anting birds. These materials include fruits, foliages, raw onion,
burning matches or tobacco, gum of grass-tree (Xanthorrhoea preissii), mil-
lipedes (Diplopoda), various beetles ("weevils"; tenebrionid beetles of the
genus Blaps ) , grasshoppers ( Anacridium aegyptium ) , earwigs ( Forjicula ) ,
bugs (Rhynchota sp., Rhaphigaster nebulosa), wasps, hair tonic, prepared
mustard, vinegar, hot chocolate, and moth balls (Ivor, 1941; Laskey, 1948;
Parks, 1945; Robinson, 1945; Thomas, 1946; Groff and Brackbill, 1946;
Baskett, 1899:24.3; Burton, 1955a, 1955b, 1955c; Chisholm, 1944, 1948:163-
175; Sedgwick, 1947; Poulsen, 1955, 1956; Sedgwick, 1946; Adlersparre,
1936; Osmaston, 1909, 1936; Callegari, 1955; Govan, 1954; Freitag, 1935;
Butler, 1910; McAtee, 1938; Fluck, 1948; Scheidler, in Stresemann, 1936;
Nice, 1952, 1955a; Hill, 1946; and others).
195
]96 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
A compilation of records shows that at least 148 species of birds, 65 of them
New World forms, are reported to introduce ants, or their so-called substi-
tutes, into or on the plumage. Included is the Wild Turkey, so far known
only for dusting in defunct ant nests (Allen, op. cit., citing Audubon; McAtee,
1947, citing Sharp). Not included are several other species, to be mentioned
later, that are known to use smoke but not ants or other substitutes.
Among the 148 anting birds listed in Table 3, the first 16 are non-passer-
ines. The Horned Owl and all but one species of the Phasianidae listed dusted
in activated ant beds or were seen to have live ants in their plmnage. The
Scaled Quail (Callipepla), the parrots, the Wryneck (Jynx torquilla) and the
Green Woodpecker (Picus viridis) applied ants or a substitute with the bill.
The belief that anting is restricted to passerines has been rather general,
and reports of anting among other groups of birds sometimes have been
questioned or discounted altogether. Inquiries and search of the literature,
however, reveal a notable amount of little known or entirely new data on a
nmnber of species, including non-passerines. Evidence for picids now is
substantial, as will be seen; and we can reasonably expect further records for
other species whose status as performers of anting still may seem suspect to
some investigators.
During my visit to Chiapas, Mexico, in July, 1956, Miguel Alvarez del Toro,
Instituto Zoo16gico del Estado, Tuxtla Gutierrez, furnished me with data on
five Mexican species he had seen anting in nature. One of these, the Golden-
fronted Woodpecker (Centurus aurifrons), used a small species of paper-
making wasp common in that region. Specimens of the wasp, received from
Dr. Alvarez in March, 1957, have been identified by K. V. Krombein, Smith-
sonian Institution, as Polybia occidentalis (Oliv.), a very gentle social species
in which only the females have a sting. Wasp venom in general is supposed
to contain formic acid.
Still another record for a picid comes from Fred M. Packard, Washington,
D.C., who advises me that he has seen anting in the Flicker (Colapres auratus)
in New Jersey (letter, July 18, 155).
Those who hold that non-passerine birds never are anting when they dig
into, and dust themselves with, ant-nest earth containing large nmnbers of
agitated, aggressive ants, may not accept Mowat's (1957) interpretation
as "ant bathing" for his tame Horned Owl's habit of "tearing an anthill apart
and then fluffing the mixture of dust and angry ants through his feathers."
Mowat (letter, April 29, 1957) states that the owl regularly engaged in the
activity on hot summer days, using the nests of unidentified, small, red lawn
ants. "He appeared to brood over these nests, after stirring them up with his
talons, and would sit for as much as half an hour without apparent move-
ment .... He showed no signs of ecstasy, or stimulation. In fact, he usually
appeared to be asleep."
Lovie M. ANTING IN BIRDS 197
Whitaker
Through the courtesy of Edwin Way Teale (1953; letter, February 6,
1955), I have the statement of Stanley Dashuta of Newark, New Jersey, who
many times has seen Ring-necked Pheasants scratch down ant hills. This ob-
server noticed hundreds of ants swarming over the birds and saw many in-
jured ants on the ground afterward.
Robert W. Darrow (Bump et al., 1947:272; letter, November 18, 1955)
found that Ruffed Grouse used both deserted and active ant nests, but in
most cases dusted in unoccupied nests or those with small populations. Never-
theless, it is by no means certain that this bird, or other birds, dusts in ant
beds only because these offer a ready supply of loose, light soil. There is
some evidence that a bird may be sensitive to special properties in the dust-
ing soil and that anting and dusting may be more closely allied than was sug-
gested by Chisholm (1948:163-175).
Howard Campbell (1954) has shown how Scaled Quail (Callipepla squa-
mata), and possibly also Gambel's Quail (Lophortyx gambelii), in four New
Mexico counties, where dusting opportunities would seem to be optimal, were
strongly attracted by places where old motor oil had been spilled. A sub-
stantial number of 46 such experimental oiled dust baths were found to have
had heavy and continuous use. In a suitable dusting area, the birds chose the
oily spots, and they worked even in an area of extremely coarse gravel which
had been oiled. Mr. Campbell wrote me that the birds seemed to use the
centers of these oiled areas rather than the edges.
At my request, he made temperature readings on treated and untreated
dusting areas, using a standard Taylor fisheries thermometer with the half-
inch bulb barely buried in the earth. A larger series of readings is needed,
but his preliminary investigation shows that temperatures of oiled soils can
be as much as 4 ø F. higher than those taken in adjacent areas.
Gibson (1954) described a White-winged Chough (Corcorax melanorham-
phus) in Australia that did not dust normally, but instead puts beakfuls of
dust into its body plumage and under the wings. The action so strongly sug-
gested antirig that Gibson, upon determining no insects were present, had
the soil tested for formic acid, with negative result.
It has been argued, largely on the basis of Walter's (1943) work, that birds
probably have little or no olfactory sense and that galliform birds, especially,
would tend to be insensitive to ant odors. But Hamrum (1953) stated the
literature on olfaction and gustation in birds is contradictory and confusing.
His own experiments show that both odor and taste probably influence food
choice in the Bob-white (Colinus virginianus). Thorpe (1956:306) cautioned
against dogmatism regarding sense of smell, pointing out that birds "show a
considerable range of development of the olfactory lobes" and that only a
few forms have been studied critically.
Until more is known about the purpose and effect in birds that apply ants
198 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
with the bill, the assumption that anting is not to be found in the galliform
or certain other groups of non-passerines (Goodwin, 1955b; Poulsen, 1956;
IJzendoorn, 1952a) seems unwarranted. At this stage of investigation, it
seems premature, even a little illogical, to say that a pheasant or a grouse,
exposing its body to a host of disturbed ants on an ant nest, is not anting but
only dust-bathing; while, at the same time, accepting as bonafide anting the
behavior of certain corvids, to be mentioned later, that neither apply ants with
the bill nor make dust-bathing motions but which stand or sit among ants
with special attitudes of wings and tail while allowing ants to invest their
plumage.
Closer study of Common Starlings (Sturnus vulgaris) and House Sparrows
(Passer domesticus) should help resolve the question of anting in game birds.
Starlings are known to work themselves deeply into the ant nest by use of the
feet, bill and wings, and to apply ants deliberately under their wings
(Floericke, 1911, and in Stresemann, 1935b; Baggaley, 1946). Davis (1945)
noted a House Sparrow on an ant hill, dusting itself among the ants; and, in
a letter to me, he emphasized that the bird also was using its bill to apply
ants under the plumage in typical anting manner.
Cases combining dusting and applying ants with the bill, such as those
cited above, are interesting also as possible transitions between "passive"
anting (standing, sitting, sprawling among ants, but usually not applying
with bill; see Fig. 5) and "active" anting (anointing only by use of bill).
This distinction is made by Rothschild and Clay (1952:126-128).
It is unfortunate that the word antirig is so deeply imbedded in the litera-
ture, since it does not have universal definition and is not always descriptive
of the behavior. Thus we find McAtee (1938, and in Chamberlain, 1954)
excluding the use of substitutes in defining anting; Goodwin (1955b) and
Poulsen (1956) dismissing records of game birds using activated ant nests--
a behavior which McAtee accepts; and Ivor (1951, 1956) excluding such
passive anting as that seen in Common Crow (Corvus brachyrhynchos). De-
spite these exclusions, it now appears that the term ought to include all anoint-
ings, whether active or passive, with ants or substitutes. Certainly this was
the connotation given the term by Stresemann (1935b) in coining it.
Although anting is a major ornithological problem that no one has ade-
quately explained, it has received relatively little experimental study. A new
theory, offered by Holger Poulsen (1955), who experimented with 34 anting
species in the Copenhagen Zoo, states that anting is caused by the ants' spray-
ing of acid on the bird's head. The bird then rubs its head under wing or
tail in an effort to remove the irritating acid. Poulsen, observing only active
anting, and apparently questioning the many descriptions of passive anting,
concluded that feeding was the basic incentive, and the anting actions inci-
dental corollaries to it, as the bird tried to cleanse its head of ant acid or
Lovie M. ANTING IN BIRDS 199
Whitaker
tripped about to avoid being sprayed. He considered tripping and falling, as
well as reports of birds lying down among ants, all to be results of the bird's
vigorous cleansing and evasive movements. Some investigators, however,
found that this theory did not explain anting as they observed it (Nice, 1955b;
Goodwin, 1955a, 1955b; Simmons, 1955; Ivor, 1956).
Since then, Poulsen (1956) continued experiments (involving 85 species
in all, of which 56 species anted) and discovered that certain species would
indeed deliberately expose their plumage to ant spray. But he still is of the
opinion that, with few exceptions, anting is unintentional behavior connected
with feeding, and that only such exceptional species (more or less passively
anting birds, as defined here) seek ants in order to be sprayed, rather than to
eat ants. While he does not use the terms, Poulsen makes it clear that he be-
lieves active anting to be a response to external stimulus and that passive ant-
ing evidently is motivated internally, depending upon the bird's being "in ant-
ing mood." He has no definite solution to the problem, and offers these ex-
planations tentatively. In a letter (February 7, 1956) he stated that he re-
garded anting as a complicated behavior, with more than one biological sig-
nificance.
In view of the many poorly understood, even puzzling, aspects of anting,
it seemed that an intensive examination of the behavior in the individual
bird, with various species of ants, might be at once interesting and worthwhile.
ACKNOWLEDGMENTS
I am greatly indebted to Dr. Robert E. Gregg, University of Colorado, for
identification of ant species, and for advice and criticism regarding myrme-
cological portions of this paper; and to Dr. Raymond A. Paynter, Jr., Har-
vard Museum of Comparative Zoology, for arranging the list of anting bird
species according to current nomenclatural usage. Dr. Poulsen kindly sent
me his papers on anting, together with some other data from his experiments.
I wish to thank Professors Eugenia K. Kaufman, .Gerhard Wiens, Joseph
Andriola and Erich H. Eichholz, all of the University of Oklahoma, for cer-
tain translations from the German and the Italian, and Mrs. Kaspar Arbenz
of Norman, Oklahoma, for translations from the Dutch. Dr. W. A. Lunk,
University of Michigan, generously assisted with some difficult bibliograph-
ical problems. I gratefully acknowledge encouragement and advice given by
Dr. Margaret M. Nice, Chicago; Mr. H. Roy Ivor, Erindale, Ontario; and
Professors George M. Sutton and Harley P. Brown, University of Oklahoma,
all of whom have seen the manuscript; and the assistance of my husband,
John R. Whitaker, in its preparation.
Special thanks are due Dr. Arthur A. Allen, Cornell University, for photo-
graphs of my Orchard Oriole in action, taken at 1/5000 of a second with
stroboscopic lighting. It was a rare and helpful privilege to be able to study
200 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
these pictures, some of which are reproduced here. Other photographs, show-
ing antirig in two European species, are used by the kind permission of Dr.
Hans L;Shrl, Ludwigsburg, Germany.
The following persons generously contributed their own unpublished ob-
servations of antirig: Miguel Alvarez del Toro, Tuxtla Gutierrez, Chiapas,
Mexico; Frank L. Beebe, Provincial Museum, Victoria, British Columbia;
Connie Hagar, Rockport, Texas; H. Roy Ivor, Erindale, Ontario; John A.
Johnson, Iron Mountain, Michigan; Levon Lee, New Mexico Dept. of Game
and Fish; Dr. G. Douglas Morris, Algoma, Ontario; Fred M. Packard, Wash-
ington, D.C.; Boris Stegmann, Zoological Museum, Leningrad, U.S. S. R.;
Edwin Way Teale, Long Island, New York; Neal A. Weber, Swarthmore,
Pennsylvania; and W. W. Worthington, Fort Lauderdale, Florida. The late
Josselyn Van Tyne, University of Michigan; Nagahisa Kuroda, Yamashina
Museum of Birds, Tokyo; Edgardo Moltoni, Museo Civicio de Storia Naturale,
Milan, were helpful in obtaining certain materials published abroad, as was
H. Roy Ivor.
AiNITIiNIG IN A CAPTIVE ORCHARD ORIOLE
On September 18, 1952, I discovered my hand-raised, three-month-old, fe-
male Orchard Oriole (Icterus spurius) antirig in a file of ants which had in-
vaded a screened porch. Systematic observations on the bird began on March
23, 1953. In the following 31 months I made 80 experiments on as many days,
using various ant species found in my yard. Excepting two experiments in
which the ant Tapinoma sessile was used in September and October, 1955,
at East Lansing, Michigan, all observations were made in Norman, Oklahoma,
with indigenous ants.
Four other hand-raised individuals--Black-billed Magpie (Pica pica hud-
sona), Loggerhead Shrike (Lanius ludovicianus), House Sparrow, Painted
Bunting (Passerina ciris)--did not ant, and only the Magpie and Painted
Bunting would eat ants. These birds were tested at intervals during periods of
two months or longer, the Magpie for over a year, while the Painted Bunting
was the constant companion of the Orchard Oriole during the latter's antirig
experiences. Kuroda (1947) noted antirig-like behavior in a captive Bull-
headed Shrike (Lanius bucephalus); but there seems to be no report of ant-
ing in Painted Buntings, and I find but three for the House Sparrow (Table
3). It is strange that there should be no New World record for this race of
Magpie, since the species is a well-known "anter" elsewhere (Table 3). In
this connection, it is interesting to recall that Brooks (1931) suggested spe-
cific rank for Pica p. budsonia.
For present purposes I shall apply the term acceptable to those ant species
the Orchard Oriole used for anointing (Table 1), and unacceptable to those
it rejected (Table 2), whether or not the species was eaten.
Lovie M. ANTING IN BIRDS 201
Whitaker
I found workers of three ant species, Dorymyrmex pyramicus, Iridomyrmex
pruinosus analis and Tapinoma sessile were acceptable and also were eaten.
Workers in these species are monomorphic. All are small (2 to 3 mm. in body
length), non-stinging ants which feed on honey-dew and insects. Instead of
spraying acid secretions when disturbed, these species exude from the anal
glands fluid droplets having the scent of rancid butter. This odor, presum-
ably due to butyric acid, is especially strong when the ants are crushed. Evi-
dently these species, all belonging to the subfamily Dolichoderinae, do not
produce formic acid. O'Rourke (1950) stated that "so far as is known, the
Formicinae [i.e., Lasius, Formica, Camponotus, etc.; see Table 1] alone
among ants secrete formic acid." For pertinent details on the ant species
mentioned above and in the next paragraph, see also Wheeler (1910:29, 42-
43, 45, 361), Creighton (1950:110-111, 162, 171, 210, 340, 346-348, 350-
352, Cole (1940), Smith (1924, 1928), Dennis (1938).
The very small ants, Pheidole bicarinata buccalis and P. b. longula, and the
small Crematogaster (Acrocoelia) laeviuscula, as well as the large Pogono-
myrmex barbatus, were all unacceptable to the oriole, although both of the
Pheidole sometimes were eaten. All the rejected ant species belong to the
subfamily Myrmicinae. Workers of these species possess a sting. The sting
in Pogonomyrmex barbatus is extremely painful to man; but that of Pheidole
is too weak to penetrate human skin, as, in my experience, was true also of
the Crematogaster. This particular Crematogaster feeds on honey-dew, dead
insects and animal tissue; but ants of the genera Pheidole and Pogonomyrmex
are largely spermophagous, though some Pheidole take insects and honey-dew,
and Pogonomyrmex will eat insect food. None of the rejected species sprays
or exudes repugnatorial liquids.
Neither the Orchard Oriole nor any of the ant species it accepted seems to
appear in antlug literature, with the exception only of Tapinoma sessile, noted
by Van Tyne (1943), and Tapinoma sp., by Ivor (1943; and in Nice, 1945).
I find no mention of butyric acid or of the fact that a non-stinging, non-spray-
ing ant species will induce anting. Thus far not enough attention has been
given the various defense mechanisms among ant species used for antlug; and
even when the ant has been identified, its particular means of defense often
has not been stated. Some writers seem to assume, quite erroneously, that all
ant species used by birds have the ability to spray or that they all produce
formic acid in quantity. Groskin (1950) believed that variations in birds'
antlug movements might be due to differences among ant species in aggres-
siveness and, he implies, in the composition of defense fluids. To this I would
add difference in amount of the repugnatorial substance and in the method
of ejaculation, i.e., whether sprayed or exuded, for reasons that will be clari-
fied later.
202 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
TABLE 1
A PHYLOGENETIC LIST OF ANT SPECIES USED BY BIRDS FOR ANTINC
Ant nomenclature found in literature on antirig is revised to conform to current usage. Insets
under specific names indicate synonyms. This informal taxonomic synonymy follows Creighton,
"Ants of N. Amer.," 1950, except for extra-limital species. Asterisk denotes species used by
Orchard Oriole.
Subfamily and Species Source
1VtyRIVtICINAE (Possess functional sting; other-
wise do not eject repungatorial sub-
stances)
Monomorium pharaonis (Linnaeus)
DOLICHODZaINaZ (Sting vestigial; exude repug-
natorial liquid from anal glands)
lridomyrmex detectus (F. Smith)
*lridomyrmex prginosus analis (Andre)
*Dorymyrmex pyramicus (Roger)
*Tapinoma sessile (Say)
FOR1VtICINAE (Sting vestigial; spray repugnatorial
liquid from special formic acid gland;
anal glands absent)
Oecophylla smaragdina (Fabricius)
Camponotus pennsylvanicus (DeGeer)
Camponotus (Myrmophyma) innexus Forel
Camponotus ( Myrmobrachys ) senex textor
Forel
Camponotus (Myrmepomis) consobrinus
(Erichson)
Lasius alienus americanus Emery
Lasius niger var. americanus
Lasius niger (Linnaeus)
Lasius niger neoniger Emery
Lasius niger vat. neoniger (Emery)
Lasius niger var. neoniger
Lasius niger
Lasius ( Dendrolasius ) /aliginosus (Latreille)
Lasius (Chthonolasius) mixtus (Nyl.)
Acanthomyops mixtus Nyl.
(Den.) Poulsen, 1955, 1956; only by
experienced Blue Jay
(Austrl.) Bourke, 1941b; Galloway,
1948
(U.S.) Whitaker, this study
(U.S.) Whitaker, this study
(U.S.) Van Tyne, 1943; Whitaker,
this study
(India) Pillai, 1941; Ali, 1936,
quoting H. Abdulali
(Can.) lvor, 1943
(U.S.) Teale, 1953:168, of "car-
penter ant"
(Austrl.) Wheeler, 1951
(Costa Rica) Skutch, 1948
(Austrl.) Chisholm, 1944; Wheeler,
1951
(U.S.) Dater, 1953
(U.S.) McAtee, 1944
(Eng.) Carpenter, 1945; Longhurst,
1949
(Switz.) Wackernagel, 1951
(Den.) Poulsen, 1955, 1956
(U.S.) Groskin, 1950
(U.S.) Brackbill, 1948
(Can. Ivor, 1943
(U.S.) Nice, 1945
(Ger.) LiShrl, 1952
(Eng.) Hobby, 1946
Lovie M.
Whitaker
ANTING IN BIRDS
2O3
TABLE 1 (Continued)
Lasius ( Chthonolasius) umbratus aphidicola
(Walsh)
Lasius umbratus mixtus aphidicola
Acanthomyops claviger (Roger)
Lasius (Acanthomyops) claviger (Roger)
Lasius clavlger
Acanthomyops interjectus (Mayr)
Lasius inter]ectus Mayr
Acanthomyops murphyi (Forel)
Lasius ( A canthomyops ) murphyi
Formica ruJa Linnaeus
Formica obscuripes Forel
Formica ru/a obscuripes Forel
Formica ruJa
Formica exsectoides Forel
Formica exsectoides exsectoides (Linn.)
Formica /usca Linnaeus
Formica usca subsericea (Say)
Formica Jusca var. subaenescens
Formica/usca s. sp. subaenescens Emery
Formica ( RaptiJormica ) sanguinea Latreille
Formica ( RaptUormica) subintegra Emery
Formica sanguinea subintegra (Emery)
Total species represented: 24.
(U.S.) Brackbill, 1948
(U.S.) Groskin, 1943, 1950
(U.S.) Davis, 1944
(U.S.) McAtee, 1938, quoting Kaita-
bach
(U.S.) Brackbill, 1948
(Ger.) McAtee, 1938, citing Heine,
1929
(Eng.) Goodwin, 1951, 1952a, 1953a
(Switz.) Wackernagel, 1951
(Den.) Poulsen, 1955, 1956
(Neth.) IJzendoorn, 1952a, citing
Abma, 1951
(U.S.) Weber, 1935
(U.S.) Nice and Ter Pelkwyk, 1940
(U.S.) Staebler, 1942
(U.S.) McAtee (in Chamberlain,
1954), "probably"; Teale, 1953:
159, 199
(U.S.) Brown, 1953; Groskin, 1949,
1950; Nero, 1951; Brackbill, 1948;
Hebard, 1949
(U.S.) Nichols, 1943, "probably
$[Lba1/21/2$C1/2l$ '
(U.S.) Buell, 1945
(Can.) Ivor, 1943, 19S6
(U.S.) Groskin, 1950
METHODS
In summer experiments, I dug up entire ant colonies (earth, ants, pupae,
larvae, eggs) and placed the material in a tray on the porch where the Or-
chard Oriole was free. Sometimes I confined the bird in a large, bottomless
cage and placed the cage over the tray, or over an ant nest in the yard.
Winter experiments were managed similarly indoors, with ants taken from
captive colonies or, occasionally, from the yard. Spraying ants, unavailable
locally, were never offered.
204 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
Each experiment, except as otherwise noted, was made with a single colony.
When more than one ant species was used in the experiment, all material of
one species was removed before another species was offered. Duration of ex-
periments varied from a few minutes to three hours, depending upon the
bird's interest and the number of ant species offered.
: ß :: : .\---
F11/2. 1. i (a) Orchard Oriole exposes undertail coverts in preparation for antirig. Note
ant in bill, and spli! web of an inner rectrix, caused by anting activity. (b) Bird ap-
parently applies ant to both rectrices and remiges, shown interlocked. (c) Application to
bases of outer rectrices. Note wing tip resting upon ground behind tail. (d) With tail
pressed against folded wing, bird treats wing tip and possibly the ventral surface of tail.
Note displaced (right) greater wing covert.
Lovie M. ANTING IN BIRDS 205
Whitaker
In three instances, two species of ants, readily distinguishable at sight, were
mixed deliberately. In 1954 most summer observations were made with ants
that daily invaded the porch in force, attracted by bird food. These invaders
invariably appeared to be all of one species, and samples from them, taken
July 30, were identified by Dr. Gregg as Dorymyrmex pyramicus.
The Orchard Oriole is trusting and permitted closest scrutiny, often per-
forming 10 inches from my face. It never has been sick or injured. Examina-
tion with hand lens and brushing out its plumage disclosed no parasites.
Since the first prenuptial molt, this female oriole has been in excellent con-
dition, with bright plumage of normal texture and color. All subsequent molts
have been autumnal. The bird's somewhat small size may be due to metabolic
disturbances as a fledgling on a partly artificial diet.
PLUMAGE AREAS ANOINTED
Although observers do not agree, and reference to application to virtually
every accessible plumage tract can be found in the literature, the wings and
tail are almost always mentioned (see Adlersparre, 1936; Allsop, 1949; Brack-
bill, 1948; Chisholm, 1944; Fletcher, 1937; Goodwin, 1953a; Groskin, 1943,
1950; Heinroth, 1911a; Ivor, 1941, 1943, 1951, 1956; Nice, 1952; Osmaston,
1909, 1936; Reynolds, 1946; Tebbutt, 1946; Van Tyne, 1943; Wheeler, 1951,
and others).
fvor (1946, 1951, 1956) modified his earlier descriptions of applications
to ventral surface of primaries by stating that ants are applied to the distal
one-third of the primaries, rather than along their entire length; that ap-
parently the undertail coverts sometimes are treated; and that the ventral
surface of the tail is treated, although rarely. Poulsen (1955, 1956), how-
ever, said that his birds, excepting the Common Grackle (Quiscalus quiscula),
applied ants only to the quill feathers.
My oriole regularly anointed the ventral surfaces of the outer few pri-
maries, mostly near the tips. In doing this, the spread tail was brought side-
ward and forward; and the wing on that side was either folded or only
slightly spread, with wrist lifted so that the wing tip was held near the basal
section of the tail, or against the ventral surfaces of the rectrices (see Frontis-
piece). Application of ants caused wing tips to become frayed and their
dorsal surfaces roughened by displacement of the barbs from beneath.
The ventral surface of the tail also was treated regularly, particularly the
basal one-third (Fig. lc). But it was the undertail coverts that seemed to
be a main target. Here the deep, vigorous applications at times caused some
of these yellow feathers to become so displaced as to stand up conspicuously
above the olive-toned upper tail coverts!
Sometimes the bird treated its belly, and very occasionally the anterior
crural feathers briefly. (For anting of the legs, see Brackbill, 1948; Hein-
2O6
THE WILSON BULLETIN
September 1957
Vol. 69, No. 3
TABLE 2
ANT SPECIES REJECTED BY ORCHARD ORIOLE FOR ANTlING
Subfamily MYRMICI]NAE (Possess functional sting; otherwise do
not eject repugnatorial substances)
Pogonomyrmex barbatus ( F. Smith)
*Pheidole bicarinata buccalis Wheeler
* Pheidole bicarinata longula Emery
* Cremato,gaster ( A cr ocoelia ) laeviuscula Mayr
* Sting does not penetrate human skin.
roth, 1911a; Nice and Ter Pelkwyk, 1940; Snyder, 1941; Tebbutt, 1946;
Troschiitz, in Stresemann, 1935b.) The flanks and sides seemed to be touched
only indirectly as the bird reached toward the wing tips and tail. The feathers
of the sides and flanks became disarranged, but I could never detect direct,
exclusive application to them. IJzendoorn (1952a), noting untidy, ruffled
breast and abdomen of a Common Starling anting in the wild at high in-
tensity, suspected the disarray indicated poor condition. The activity itself
caused somewhat the same appearance at times in my oriole.
In the rare instances when the oriole applied ant larvae and pupae, it
usually ate them afterward. Sometimes a larva or pupa was used when the
bird applied the ant which carried it. On some days the bird ate directly
eggs, larvae and pupae, yet often these were ignored. Burton (1955a) ob-
served anointing with ant cocoons, the bird being a tame Rook (Corvus ]ru-
gilegus), which probably "at some time picked up an ant carrying a cocoon
and now associates the two."
On a number of days I examined the oriole immediately after anting.
found ant odor on all these areas mentioned, but no odor on other plumage
parts. The odor was always strongest on the undertail coverts and adjoin-
ing portions of rectrices. For example, on February 24, 1955, ! tested the
bird after it had anted at high intensity for 18 minutes with freshly dug ants
and ! found odor on tips of the primaries, on undertail coverts, basal two-
thirds of the tail, and on sides and flanks. The feathers of the fronts of the
tibiae were faintly scented. Odor was strongest on the undertail coverts.
Save sides and flanks, ! had clearly seen the bird treat all these areas. I
could detect no odor on other plumage areas and none on the wings, except
on the distal one and one-half inches of the primaries. These were, be it re-
membered, non-spraying ants. Neither on this occasion nor on any other
have ! seen the bird apply ants to dorsal surfaces of wings, tail or body.
! find but three references to odorous plumage, all relating to birds that
used spraying ants. Wackernagel (1951) said his tame Carrion Crow
(Corvus corone) became so scented with Formica ru/a that it was still odor-
ous next day. The scent of Lasius ]uliginosus on tame European Jays (Gar-
rulus glandarius) was evident at a meter's distance (Ltihrl, 1952, 1956). Mr.
Lovie M. ANTING IN BIRDS 207
Whitaker
John A. Johnson (letter, Sept. 17, 1954), in Michigan, informs me that his
tame, free-flying Common Crow, which has sought unidentified ants of its
own volition during nine summers, subsequently reeks with odor to the de-
gree that his own hands become scented from handling the bird. From his
description, I judge this bird used spraying ant species. All spraying ants
belong to the subfamily Formicinae (Table 1).
POSTURING, TRIPPING AND FALLING
The awkward posturing and strange acrobatics of the oriole were similar
to those described for many small species. Always there was some deflection
of the tail, even during low intensity anting. Usually the tail was brought
sharply around to one side of the feet, or between them, so that the tip
pointed forward and the dorsal surface lay largely on the floor. Often the
bird stood on the tail with one or both feet, or briefly sat on it. It habitually
tripped on its tail, at times falling sideward or backward. Sometimes it
turned complete [orward rolls. Ivor, Poulsen, Van Tyne, Groskin (1950),
Adlersparre, Osmaston (1909), Nichols (1943), the Shackletons, Nice (1943),
LiShrl (1956), and others have observed falling or tumbling in anting birds.
The higher its anting intensity, the more the oriole tripped and tumbled.
I have seen it stand with the tail turned out to one side while it looked for
the next ant; but usually it kept these odd positions only momentarily, and
after tripping or falling, would right itself in a flash. All imbalance oc-
curred in conjunction with interference of the tail with feet. Never did this
bird lie down, press its breast to floor, sprawl with spread wings, or behave
in any way that suggested passive anting.
Likewise, the oriole never permitted ants to crawl upon it, and would
quickly pick off those that got on its toes, often flipping away others that
came too close. There was no evidence that the bird ever deliberately de-
posited ants in or on the plumage. Three times only did I see single ants
sticking to its plumage, on a rectrix. Examinations in the hand revealed no
ants on the bird. However, the projected transparencies show that ants some-
times were rubbed off on the feathers. In one picture two ants may be dis-
tinguished on the ventral surface of rectrices, in another an ant is seen on the
ventral surface of a primary. These ants appear to be flexed; one of them
plainly is wedged between barbs.
The literature indicates that only a few small birds, such as Redwing
(Turdus musicus), Song Thrush (T. philome'los), European Blackbird (T.
merula), American Robin (T. migratorius), Catbird (Dumetella carolinen-
sis), Common Starling, and Indigo Bunting (Passerina cyanea), sometimes
allow ants to crawl upon them in numbers (Bates, 1937; Callegari, 1955;
Floericke, 1911, and in Stresemann, 1935b; Groskin, 1950; Govan, 1954;
Shackleton and Shackleton, 1947; Poulsen, 1956).
208 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
MANNER OF APPLYING ANTS
Observers often mention a stroking or preening action of the bill as ants
are applied (Adlersparre; Goodwin, 1955b; Brackbill, 1948; Buell, 1945;
Van Tyne, 1943; Ivor, 1943, 1956; Laskey, 1949; Wheeler, 1951, and others).
This was not the case with the captive Orchard Oriole. It always applied ants
in quick dabbings, rather than strokings. Moreover, there was also a pe-
culiar motion of the head. As the bird, ant in bill, reached toward the area
to be anointed, it moved its head from side to side, so rapidly that the ef-
fect was almost that of shuddering. While reaching for the plumage and
during each dabbing, thrusting application, the bird vibrated its head in this
manner. When performing in a clinging position on the cage wall, the oriole
sometimes caught its bill in the hardware cloth as it tried to reach its tail,
and the bill rattled loudly and rapidly between the wires. This action on
quill feathers produced a rustling sound. Invariably ants were applied thus;
but during low intensity anting, vibration and dabbing were likely to be
shorter in duration and noticeably slower.
Though these head movements have not been described precisely here-
tofore, they probably occur in many birds. Adlersparre said only that two
Orange Bishops (Euplectes franciscana) shook their heads so energetically
they sometimes lost the ant from their bills. Poulsen (1956) stated that his
birds would "rub the bill among the wing-feathers downward towards the
tip with quivering movements of the head"; and he mentioned that they
sometimes would shake their heads "more or less vigorously." Both of these
writers seem to attribute this behavior to ant spray, a factor not present in
the case of the oriole. In response to my query, lvor (letter, February 16,
1955) wrote that he noticed, on that day, both the dabbing applications and
vibrating head for the first time, in the case of a Baltimore Oriole (Icterus
galbul); and, further, that he believes that, in working with groups of ant-
ing birds, he had previously overlooked these motions. Bourke (1941b) told
of Rufous Whistlers (Pachycephala rufiventris) that took ants to bushes and
there shook their heads from side to side a few times, then dropped the ants
and preened. Head shaking preceding ant applications has been reported in
Indian Mynahs (Acridotheres tristis), according to Chisholm (1944); and
the Cockatoo observed by Glauert behaved similarly with an ant and again
when tobacco juice got in its mouth.
Among numerous responses lvor received, following his recent paper
(1956), was one from Henry Petersilie in New York, suggesting that birds
may fall over in "ecstasy" because of disturbance in the semicircular canals
of the inner ear. The idea has merit. Granted a disturbance to balance mech-
anism, from head shaking or other cause this alone would not seem enough
to have caused the oriole's loss of footing; for the bird was not seen to trip
or fall except when the tail was brought into contact with feet and legs. But
Lovle i. ANTING IN BIRDS 209
Whitaker
there remains the possibility that loss of balance through interference of
the tail was augmented by some such condition.
TREATMENT OF ANTS
The captive Orchard Oriole seemed to roll these small ants excessively.
Usually each one was worked in the bill, and apparently crushed, regardless
of its ultimate disposition. Supplies of acceptable dead ants were treated in
like manner. I could not tell how severely the bird damaged an ant before
applying it; but ants that it had rolled and cast aside unused were so injured
they could not crawl. Those flung away after application were either dead
or severely injured. Examination of 14 such "used" ants under binocular
microscope showed 13 with damaged gaster, several with tissues extruded.
Some of them also had the pedicel, head or thorax injured and nearly all had
lost appendages.
The bird habitually made several applications, each consisting of many
tremulous dabs, with a single ant. Between applications, it would hesitate
and roll the ant further. It seemed to me this was done in order to renew or
increase the ant's effectiveness.
A number of records show squeezing or crushing of the ants (Troschiitz,
in Stresemann, 1935b; Ali, 1936; Nice and Ter Pelkwyk, 194,0; Galloway,
1948; Poulsen, 1955, 1956; Teale, 1953:168) or that dead and injured ants
were left on the anting grounds. Ivor (letters, February 1 and 14, 1955) and
Teale (loc. cit.) each have noticed that workers of the large spraying ant
(Camponotus pennsylvanicus) sometimes seem not to be crushed before be-
ing applied. Teale, Groskin (1950) and others have suggested that spraying
ants may give off acid merely by being held in the plumage. Perhaps a bird's
treatment of the ant may depend more upon the ant's spraying ability than
upon its size. Formica rufa and its allies can eject a fine spray for a distance
of 20 to 50 cm. (Wheeler, 1910:42-43). It should be recalled that my
oriole used only non-spraying ants, and that crushing increased their odor.
I found, however, in at least one of these species (Dorymyrmex pyramicus),
that only the gaster seemed to produce odor. When the gaster was removed,
crushing of the head, thorax and other parts did not seem to change or in-
crease the ant's odor.
THE ANTING LOCALE
Birds have been observed anting in trees, on roofs or feeding tables--
wherever ants happened to be found. Sometimes they carry ants from ground
to a tree or bush to apply them, whereas a captive bird may take them to
its perch.
Usually my bird anted on the floor. Sometimes it performed on a perch,
either bringing up ants or using those that crawled within reach. But it also
had a habit of anting whil clinging to the cage wall. It might fly to the
210 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
wall with an ant and cling there to apply it, or hitch its way down the wall,
snatch and apply an ant without touching the floor. At times the bird anted
while clinging upside down on the wall--a position especially favorable for
observing anointment of belly or undertail coverts. Skutch (1948) saw a
Black Seed-eater (SporophiIa aurita), anting in a bush, make one application
while hanging upside down from a twig!
ATTACKS BY ANTS
Surprisingly few accounts show that anting birds are attacked by the ants
or that they fear them, although some kinds used are formidable biters.
Aside from Poulsen's Blue Jays (Cyanocitta cristata), which used a small
species (Monomorum pharaons), it appears that birds use only those ants
incapable of stinging (Tables I and 2). Furthermore, Poulsen (1955, 1956)
thought the Blue Jays' responses were due to conditioning. His birds in
general rarely were cautions or hesitant in taking up ants; and, when bitten,
would shake their legs, jump or pick off the ant and fling it away.
Groskin (1950) saw a Song Sparrow (Melospiza melodia) suddenly jump
and dance around before resuming anting; Ivor (1943) said "numerous
times it was evident an ant had bitten a bird"; Bourke (1941b) commented
on the hastiness with which Rufous Whistlers removed ants gripping their
feet; Wackernagel's Carrion Crow at times "showed severe fright reactions
and jumped into the air, probably when bitten"; and LShrl (1952, 1956)
described similar behavior for this crow. Goodwin (1952a) noted fear and
hesitation in Lanceolated Jay (Garrulus lanceoIatus) and Beechey's Jay
(Cssolopha beecheli) but not in some other species which apparently used
the same ant species. These jays would first take ants to perches to apply
them before anting on the ground near the ants.
My oriole approached the unacceptable Pogonomyrmex barbatus readily,
yet carefully; but it showed no fear of other ants, except, as will be seen, with
an acceptable species foraging en masse. True, throughout the period of
study, the bird performed at times on wall or perch; but it seemed to fly up
with an ant in quite the same way it carried off a grasshopper to be eaten.
Moreover, with supplies of dead ants the bird behaved this way. Although
it would pick up unacceptable ants, even eat certain species of them, I never
saw it carry an unacceptable species from the floor.
On June 30, 1954, I discovered a file of ants reaching from the porch door
to a screened section occupied by the Magpie. The ants came in force almost
daily until September 30, and during this period I put the oriole with them
at least once a week. As previously mentioned, probably all of these ants
were Dorymyrmex pyramicus.
On July 8, after eight successive days of antlng with these invaders, the
oriole began performing mostly on perch or all. In the following three
LovieM. ANTING IN BIRDS 211
Whitaker
weeks, it showed increasing reluctance to perform on the floor. On July 12,
16, and 24, all anting took place above the floor. Yet, on July 31 and August
1, the bird anted only on the floor. This erratic behavior strongly suggested
fear of attack. Probably the bird had been bitten at one of the times when
it had been allowed to remain with ants after the observation period. Later,
on August 3, I witnessed an attack that was to affect its performance for quite
some time.
As I watched the oriole ant, it suddenly jumped straight upward and then
began to probe between a middle and outer toe. Almost at once it flew to
a perch and began probing gently at the spot. For several minutes it re-
mained there quietly, looking somewhat subdued, now and then peering down
at its toes. Presently it began performing with ants taken from the wall; but
not again that day did it go to the floor, except once to snatch an ant and
fly with it to a perch. During the following two weeks ! tried the oriole with
invading ants on seven days. Each day the bird anted, but only above the
floor, although on each of two days it did perform a single anting on the floor.
GATHERING AND APPLYING WADS OF ANTS
Sometimes the oriole gathered a ball of ants in the bill tip and held it for
a while before eating the wad, applying it, or flinging it away. Once, when
anting interest was very low, the bird held a wad of ants for almost two min-
utes while sitting still on a perch. One or several wads were gathered on 14
days, including experiments with dead ants. I once counted 18 ants taken
up in rapid succession. Some wads were larger. The ants, clinging to one
another like filings to a magnet, were so injured they did not separate after
being cast away. When a wad was applied, parts of it dropped off and the
bird seemed to eat the remainder. Of unacceptable ants, only the Pheidole
were gathered, and then eaten.
Not very many anting species have been reported to gather ants. The Blue
Jay, observed by Buell (1945), evidently applied small masses of ants.
Gough (1947) watched Song Thrushes apply one or two ants at a time but
not the large numbers which some of the birds collected. Ringleben (in
Stresemann, 1935b) did not make clear whether his Carrion Crow made use
of the wad of ants before throwing it aside. Scheidler (in Stresemann, 1936)
and Ivor (1956) mentioned captive Common Starlings filling their bills with
ants and then rubbing them on the plumage. Gengler's (1925) captive Com-
mon Starlings and Funke's (1912; and in Stresemann, 1935b) tame Magpie
used several ants at a time. Simmons refers to this behavior in these last
two species, as do Goodwin (1955a; 1955b) and L/3hrl (1956) to the Star-
ling. Poulsen (1956) said that the Blue Jay and Common Starling often re-
tained ants after applying them and that, by the process of applying and
then retaining ants, these birds collected as many as 20 ants before discard-
212 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
ing or swallowing the mass. On one occasion he saw this in the American
Robin.
REACTION TO ANT-NEsT EARTH
A strange activity of the Orchard Oriole concerned not the ants them-
selves but their nests. On seven days, winter and summer, the bird ate par-
ticular bits of nest earth, taking soil only from one or two minute spots in
the tray. It always seemed to examine the dirt very closely before finding a
spot to its liking. At times I could see what appeared to be grains of earth
on its long tongue. Once, after several daily anting sessions, the bird was
surfeited and would not ant--yet it ate a little of the nest earth. On two
days, with the unacceptable Pheidole bicarinata buccalis and Pogonomyrmex
barbatus, it ate nest earth. Ants, eggs, larvae, pupae were not involved in
these feedings, though the bird may have been finding infinitesimal pellets
of food residue ejected from the ants' infrabuccal pockets or, conceivably,
minute myrmecophilus arthropods (Wheeler, 1910:32, 378-397).
On February 24, 1955, I tested the bird with earth (free of ants) from a
nest in the formicarium. The oriole ate certain bits of the earth; yet when
I replaced this with garden dirt, the bird hunted over the tray but did not
once touch bill to the soil. Next I gave it still another sample of formicarium
earth. Again the bird ate soil particles. Twenty minutes later it anted at
high intensity with ants fresh from the yard, but it ate no dirt.
I have seen no record of such feeding. Gravel was always available to the
bird; and powdery soil, which it spurned for dust-bathing, it never ate. It is
a guess that the oriole ate only soil particles impregnated with ant exudations,
possibly soil that had lined brood chambers. Wheeler (1910:395) stated
that galleries of populous ant colonies becomes "greasy from the attrition of
the constantly passing ants." Ivor (letter, May 10, 1956) suggested that the
oriole may have found some kind of beneficial mold in the nest earth.
FREQUENCY OF ANTING EPISODEs
Little is known concerning frequency of anting in the individual bird.
Groskin (1950) observed a banded Song Sparrow, probably two individuals,
anting on five days during one month, often "several times a day"; and Mayr
(1948) saw a Song Sparrow ant almost daily in July in a certain stand of
dock (Rumex) frequented by ants. Three Indigo Buntings in the wild used
ants on four consecutive days (Shackleton and Shackleton, 1947). John A.
Johnson (letter, September 17, 1954) advised me that his tame Common
Crow ants perhaps once a month or whenever the weather is warm and dry.
Goodwin (1951) found his six European Jays would not ant "two or three
days running" and that a week or more seemed necessary between sessions
for "keen" performance. Ivor (1951:177) believed his birds would ant only
occasionally if ants were kept in the aviary.
Lovie M. ANTING IN BIRDS 213
Whitaker
On the other hand, Poulsen (1956) stated: "Many birds eating and anting
with ants did so every day--sometimes several times daily--for more than
a month." But he found that some of the species which applied ants but also
allowed ants to invade their plumage (Turdus philomelos and T. musicus)
did not respond again until three days later, while an American Robin, hav-
ing refused to ant for three consecutive days, anted on the fourth day.
Study of individual differences in amount of anting is long overdue. Why,
among captive Magpies (Pica pica), should one bird ant and the others never
ant? Why did Goodwin's (1955b) Lanceolated Jays ant at their first op-
portunity, in 1951, and thereafter refuse? Why did some of Ivor's (letter,
November 25, 1955) anting birds, among them individuals that had anted
for years, either refuse or ant only sparingly during repeated tests in spring,
summer and fall of 19557
These and most references to frequency of anting or to absence of anting
in experimental birds are unsatisfactory in that there are no precise state-
ments on the quantities of ants supplied, and often the ants have not been
identified. Poulsen (1956) gave his birds "a shovelful of earth containing
several ants." Sometimes my oriole ate many ants before beginning to apply
them. Thus, a bird given a limited daily supply might be expected to show
a different frequency of anting from that occurring when its supply is abun-
dant enough to permit daily surfeit. I believe that the amount of insect food
in the diet of a captive bird also may affect anting responses, and that a cap-
tive starved for fresh insects may eat all of a limited supply of ants without
performing any anting.
In this connection, I should emphasize that whenever the oriole had access
to invading ants (see Methods) these were in almost limitless supply and
continued to invade long after the bird had lost all interest in them. Also,
in a majority of the other experiments more ants were offered than the bird
could use.
I found anting frequency was high in my oriole, at least in summer. In
1954, between June 30 and September 19, the bird was put with invading
ants on 41 days. Of these 41 days when there was opportunity to ant with
the same species of ants in their natural state, the bird performed on 34 days.
On three other days there was circumstantial evidence of anting; for, on
these days, I did not remain to watch the bird, yet later I found scores of
dead ants on the floor. On only four days did the bird refuse to ant.
In July alone, during 24 contact days with these ants, the oriole anted on
19, gave circumstantial evidence on three others (the three days mentioned
above), and refused to ant on only two days. Also, on three of the 19 days
(July 5, 8, 9) the bird was placed with the ants both morning and afternoon,
and it anted each time.
The longest period of successive daily performance by the oriole was 10
214 THE WILSON BULLETIN Septelnber 1957
Vol. 69, No. 3
days (June 30 to July 9); the next longest was seven days (July 31 to August
6). Both sequences occurred with invading ants.
For the entire 31-month study period (March 23, 1953, to October 6,
1955), negative response was even lower. Out of 77 contact days (not in-
cluding the three days of circumstantial anting), the oriole's response was
positive on 67 days and negative on 10 days. Analysis of the 10 refusals
shows that the bird refused because of apparent surfeit on only four days.
On the other six days rejection was due either to the presence of strangers
(one day, with acceptable ants available) or to the fact that only an unac-
ceptable ant species had been offered (five days).
DURATION OF ANTING EPISODES
Anting birds frequently perform from a few minutes to one-half hour.
Two Scarlet Tanagers (Piranga olivacea) anted, with short interruptions, for
more than one hour (Groskin, 1943); three American Robins anted in turn
for over 45 minutes (Nichols, 1943); and a Common Grackle used green
fruits of Magnolia acuminata for over an hour (Parks, 1945). In the case
of three Cardinals (Cardinalis cardinalis) observed by Kurata (Snyder, 1941)
to ant for about two and one-half hours, it is not clear whether the birds were
under continuous observation or that this constituted a single episode.
My oriole commonly anted for 25 minutes; that is, from the time it be-
gan to apply ants until the time it quit or the ants were removed. Frequently
it performed for 45 minutes and was still anting when I ended the experi-
ment. On May 22, 1954, the bird anted for 19 minutes with dead ants, and
soon thereafter, for 30 minutes with live ants. Again, on April 3, 1954, during
one hour and 55 minutes, when the bird was given three separate and vary-
ing supplies of ants, it performed for 3, 13 and 20 minutes, in that order.
Winter sessions at times were equally long. On January 14, 1955, the oriole
anted intermittently for about 90 minutes while Dr. A. A. Allen was taking
pictures. Two weeks later it anted for 45 minutes, when there had been no
unusual distractions.
As a rule anting continued quite steadily, with brief intervals now and
then when the bird might search for ants, rest, or engage in unrelated ac-
tivities. When anting at top intensity, the bird applied ants, one after the
other, as fast as it could snatch and use them. But after a few minutes of such
rapid action, it would stop anting and stand still for a while, as if exhausted,
before resuming. I was impressed by the bird's strenuous, apparently com-
pulsive, exertion. During warm weather, in strong sunshine, it sometimes
stood and panted, but its non-anting companion, the Painted Bunting, did
not react in this manner. After long, vigorous sessions, the oriole would feed
and then sit quietly for as long as 45 minutes, sometimes sleeping. Ringleben
(in Stresemann, 1935b), who stated that his Carrion Crow obviously was
Lovie M.
Whitaker
ANTING IN BIRDS
215
FIe. 2. (a) Orchard Oriole, resting after anting, reveals displaced plumage of belly
and side. (b) With one ant wedged in the left outer rectrix (showing below inner toe),
bird reaches for another ant. Ic) Oriole, ant in bill, almost doubles body in reaching
toward juxtaposed wing and tail. Note open eyes. (d) Bird resumes normal pose after
applying ant, still held in bill. Note lumps of earth thrown upward by motion of tail.
216 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
tired after anting for about 25 minutes, appears to be the only other observer
who has noted fatigue.
SELECTIVITY IN THE ANTING BIRD
Goodwin (1951:621-623) said of captive European Jays, regularly anting
with, but never actually applying, Formica ru/a: "Attempts to induce ant-
ing with small numbers of other species of ants have failed, possibly because
they were not offered in sufficient quantity." But he does not name the ant
species. I found that the oriole would respond to few ants or to one ant.
Adlersparre (1936), Nice (1943, 1955a), Nice and Ter Pelkwyk (1940), all
had the same experience with actively anting birds of several species. Thus
it seems that release of anting response in birds that apply ants is not de-
pendent upon the quantity of ants. Certainly my bird consistently rejected
abundant but unacceptable ant species, including harmless ones which it
often ate. But I believe the quantity of ants can affect intensity of the per-
formance. Anting in the oriole began at low intensity, as a rule, soon built
to a peak, and continued in long plateau before gradually subsiding.
I never saw my bird use a substitute. Daily it fed on apple and orange,
both of which are known substitutes (Hampe, in Stresemann, 1935b; Chis-
holm, 1944; Laskey, 1948; Nice, 1952). Hampe (loc. cit.) and Poulsen
(1955) both reported that vinegar induced anting. But when I once put
vinegar solution in the oriole's honey-water vial, the bird seemed puzzled.
Three times it plunged the bill into the liquid, backing off as if in surprise
and shaking its head. This bird also avoided burning cigarettes and fled
their smoke. Fifteen wiggling, inch-long larval centipedes (Chilopoda) ex-
cited the bird but it would not approach them. After I had crushed the heads
of several, the oriole took one but soon tossed it away.
Certain species, even individual birds, have used ants and one or more
substitutes (Adlersparre, 1936; Poulsen, 1955, 1956; Burton, 1955a, 1955b;
Nice, 1955a; Nice and Ter Pelkwyk, 1940; Scheidler, in Stresemann, 1936;
Alvarez del Toro, MS). Dr. Alvarez found the Streak-backed Oriole (Icterus
pustulatus) in nature using a species of small wasp, as well as two species
of ants. Burton's astonishing Rook used ants, both burning and hot (but
extinguished) cigarettes and matches, and small live embers, besides perform-
ing the same movements of anting whenever it could get within reach of
smoke. This Rook, according to its former owner, reacted similarly to steam
from an electric kettle, even knocking off the lid at times in order to reach
the steam. A number of times it performed in front of an electric heater, ap-
parently stimulated by the heat. But Burton's European Jay, though re-
sponding to snubbed but still warm cigarettes or blown-out but smoking
matches, refused to use the unidentified ants, cold tobacco and the several
other common substitutes offered it.
LovieM. ANTING IN BIRDS 217
Whitaker
Dr. P. H. Fluck (1948), whose tame Blue Jay used various bitter, sour
fruit juices and hair tonic, informed me (letter, March 24, 1956) that he has
a second such bird that anoints only with the hair tonic.
Poulsen (1956) noticed that anting in the Magpie-Robin (Copsychus sau-
laris), Shama Thrush (C. malabaricus), Peking Robin (Leiothrix lutea), and
Common Starling was more pronounced with Formica ru]a than with Lasius
niger. He pointed out that this last ant is smaller, weaker in its bite, and
produces less spray. His Western White-Eyes (Zosterops palpebrosa) and
Blue Sugarbirds (Dacnis cayana), on the contrary, were much more likely
to perform with this smaller ant and were cautious with the larger species
and rarely used it. In his opinion, all of this appeared to indicate that bird
species differ in their sensitivity to the bite and spray of ants. One might
mention that this also was a case of the smaller birds showing preference
for the smaller of two spraying ant species.
My exploratory experiments with selectivity in the oriole yielded some
interesting resuks. Whenever I offered either of the Pheidole forms, the bird
usually ate them sporadically but never applied any of them. Yet immediately
thereafter, it would ant at length with Dorymyrmex pyramicus. At the very
first experience with Pogonomyrmex barbatus, the oriole seemed to sense
its harmfulness. Always the bird handled this species gingerly, taking up
the ant with a pick-flick motion that sent it tossing. When repeated treat-
ment had stunned the ant sufficiently, the bird would pinch it slowly a few
times before discarding it. On one occasion pinching either brought out a
distasteful flavor, or the bird was stung; because suddenly the bird flung
the ant aside and began working its tongue, shaking its head and repeatedly
wiping the bill on the window sill where it had been standing. Never did
the oriole eat or apply this species, although immediately afterward it would
do so with acceptable ant species.
The ant Crematogaster laeviuscula was completely ignored. Indeed the only
time I saw the bird touch an ant of this species was when ! offered them from
the hand and then they were thrown away at once. In the summer of 1955,
this species, not Dorymyrmex pyramicus as in the previous summer, invaded
the porch almost daily from mid-July to mid-August. The oriole was put
with these ants a number of times; but, as far as I could determine, this
species was rejected, both initially and repeatedly, without as much as an
incipient peck. Since the bird seemed to enjoy killing the aggressive Pogono-
myrmex, whose sting and bite both presented threats, it seems unlikely that
the weak sting of Crematogaster was the deterrent. In any case, neither of
these two ants is particularly odorous, even when crushed.
In addition to the four above-mentioned ants, the oriole also rejected a
color variety of an otherwise acceptable species. This happened with Dory-
myrmex pyramicus, an ant that occurred in my yard in two color varieties,
218 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
one blackish, the other pale amber. On June 21, 1954, ! offered a nest of
the amber-colored ants. The oriole looked them over casually but would not
touch them, or the larvae and pupae, during 15 minutes. It did, however,
once eat a bit of the nest-earth. I then added a nest (no larvae or pupae) of
the readily distinguishable dark-colored ants. As I poured out these ants,
the oriole was instantly alert, crouching and leaning out on its perch, as if
in interested recognition. As soon as my hands were out of the way, it be-
gan to ant. But not once, during about 20 minutes of antlug, did I see the
bird take up a light-colored ant. Each time it chose a dark ant from the
mixed colonies. (Specimens from both colonies, taken at the time, were
identified by Dr. Gregg.) Three days later I brought in from the original
nest site the remnant population of light-colored ants. Again, the bird re-
fused to touch them. Regrettably, I did not test flavor and odor in these ants,
and I never again found this pale variety.
Was rejection due to color? Or might the pale ants have been callows and
thus less strong in odor and related qualities? According to Wheeler (1910:
534), the young worker ant first develops its own individual odor during the
period when the integument is hardening and taking on adult coloration.
Fielde (1905) has shown that a worker ant's individual odor intensifies or
changes with age "to such a degree that they may be said to attain a new
odor every two or three months" and that hostility between colonies of the
same species and variety may be caused by a difference in odor "coincident
with difference in the age of the colonies." Morley (1941-1942) said that
Fielde's work seems to show that ant odor is not fully developed until some
time after callow stage and the ant is fully adult. Not recognizing at the
time that age of ants might be of possible significance in anting, I did not
preserve the pale specimens, once they had been identified.
A thorough study of selectivity needs to be made. No one knows pre-
cisely how or why birds make their choices of ant species or substitutes, or
whether conditioning on a particular anting material actually occurs. Various
non-anting responses of birds, some of them known antlug species, to wasps
(Hindwood, 1955; Goodwin, 1952b; Chisholm, 1952; Moreau, 1942; Rankin,
1950; Powne et al., 1951), together with other cases of definite anting with
wasps (Alvarez del Toro, MS; Freitag, 1935; Butler, 1910), suggest that
birds may be similarly sensitive to differences in wasp species, in some of
which both sexes are stingless. Butler (op. cit.) said, of Garrulax sp., only
that "the wasp is seized and its tail rubbed backwards and forwards between
the tail-feathers of the bird, either in order to break the sting or exhaust its
venom before it is eaten."
REACTION TO DEAD ANTS
Both stunned carpenter ants (Camponotus sp.) and dormant ones have
been applied by various small birds (Nice, 1943:81; Ivor, 1943, 1956). Of
Lovie M. ANTING IN BIRDS 219
Whitaker
dead "wood ants" (apparently some of them killed in test tubes in boiling
water), Adlersparre said only that these ants caused his birds to perform
as if with living ones.
On April 3, 1954, I placed 40 Dorymyrmex pyramicus on a metal tray,
passed it momentarily over low heat to kill the ants, and offered it to the
oriole. The bird came at once and ate two ants. Its manner was casual and,
during 10 minutes, it applied but four ants, two of which were afterward
eaten, and ate 10 others directly. The dabbing and vibrating movements of
the bill were very much slower than usual. There was no tripping, and only
the wing tips were treated. The actual anting span was about three minutes.
I then offered approximately one-half of the nest from which these ants
had been taken, together with its living workers and larvae, but no pupae.
Now the oriole anted for 13 minutes, using all ants that did not crawl out
of reach. Anting intensity was high. Many ants were eaten, with and without
application. Forty minutes later, I replenished the tray with the remaining
portion of the nest. This time the bird anted at top intensity for 20 minutes,
after which I removed the ants. It rolled, fell down and tripped; but it ate
few ants and incapacitated ones littered the floor. The usual plumage areas
were anointed; and I noted three applications to the crural tracts.
Substantially this same experiment was repeated seven weeks later, on May
22, when I offered first 50 heat-killed ants (probably the Dorymyrmex), and
then their nest containing live ants and pupae. On this day response to heat-
killed ants was somewhat more pronounced. The bird performed intermit-
tently with dead ants for 19 minutes, but with a marked lack of eagerness.
Again movements were slow, yet at least 13 applications were made to basal
section of the tail (once to undertail coverts alone) and eight to wing tips.
Eight dead ants were eaten before antirig began. Most ants applied were
afterward eaten, but I noted nine others were eaten directly. Again, live ants
induced intense anting until they were removed after 30 minutes.
A third such experiment next day, with Iridomyrmex pruinosus analis,
brought a different result. When 150 heat-killed specimens, together with
eggs but no larvae or pupae, were offered, the bird refused to perform dur-
ing 17 minutes. In the first 11 minutes the oriole expressed some interest
and ate nine ants; then it hunted over the tray and floor as if seeking live
ants, now and then probing at the dead ones without picking up any.
Ten minutes later, I tried the bird with 150 live ants from the same colony,
placing them on the bare tray, along with a few eggs. Instantly the oriole
was attentive. It ate three ants and then began to perform as usual but at
moderate intensity, until only a few dead or maimed ants remained. Twenty-
four minutes later I gave the bird the nest of this same colony with the re-
mainder of its population, including eggs, larvae and pupae. The oriole ate
four ants, then anted at moderate intensity for 15 minutes.
220 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
On November 1, 1954, the oriole applied and ate the few dead, wet ants
(probably the Dorymyrmex) offered it. These ants had drowned in a formi-
carium moat. Next day I offered on a saucer about one-half teaspoonful of
drowned ants which had been dried at room temperature. There was quick
interest, but the bird, unfamiliar with the saucer, seemed afraid. Twice it
climbed down the cage wall until it could reach an ant, then applied the ant
while clinging upside down. Thereafter the oriole anted readily on saucer
and floor for 12 minutes. The usual areas were anointed, including the under-
tail coverts. When all ants had been "used," the bird hunted for more and
applied discarded ants, some of which had been applied previously. Anting
intensity was high. I counted 55 ants applied, some of which were afterward
eaten, and a number eaten without previous application.
Frozen ants (probably the Dorymyrmex) from several colonies, collected
in early November, 1954, and stored in a tight but not air-tight container,
were also used by the oriole many times that winter through January 18,
1955. I had drowned, rinsed and drained the ants before freezing them.
Whether few were offered or many dozens, the bird always performed. Given
small numbers, the oriole usually ate each one, about half of them being first
applied, and often it hunted for more. It used frozen ants with somewhat
less enthusiasm than it exhibited with live ones, yet the usual plumage tracts
were treated and action and interest usually were keen.
Then, on January 31, 1955, the bird ate 16 of approximately 100 frozen
ants but it refused to ant during 13 minutes. Five ninutes later it gave an
average performance with live ants from the formicarium. By this time the
frozen ants, stored over two months, had lost much of their odor and flavor,
a significant change to be described later. Furthermore, frozen ants there-
after were eaten but never again applied, although the bird continued to
perform with living ants.
REACTIONS TO SUBTLE VARIATIONS IN THE ACCEPTABLE ANT
Each ant species accepted by the Oriole has only a simple worker caste; and
sexual forms never were offered the bird. Nevertheless, during anting ses-
sions, many ants were rolled and flung aside without being applied. Oc-
casionally the bird would then toss its head and wipe its bill as if the ant
had been mildly disagreeable. But in the next moment, another ant of the
same species and colony would be rolled and applied, while a third one might
be rolled and dropped. Frequently the bird returned to such discarded ants,
even wads of ants, picked them up, rolled and discarded them again, still
without applying them.
It was Margaret Nice who stimulated ne to taste ants. ! crushed between
the teeth and tasted many ants from the supplies given the oriole, testing
every species offered.
Lovie M. ANTING IN BIRDS 221
lrhitaker
I found all of the acceptable species, fresh from the yard, usually were sweet
at first and then they produced a mildly burning sensation on the tongue.
Strangely, individuals of a colony varied a few of them, although sweet,
did not burn. I found also that odor among these ants varied. Usually they
were strong-smelling when crushed between thumb and finger, but some in-
dividuals gave off little or no odor. Larvae and pupae were faintly sweet,
but did not burn the tongue; nor did these have the bntyric acid odor so
characteristic of adult workers. (According to medical science, strongly ir-
ritating substances, such as mustard or formic acid, applied to the skin, can
produce measurable amounts of heat, due to dilation of the capillaries. There-
FxG. 3. Greatly enlarged section of Fig. 2c, showing effects of vigorous anting in Or-
chard Oriole. Note (a) rents in four outer primaries caused by bill; (b) ant-nest earth
(showing as white specks in photo) on bill just to left of ant, and adhering to edges of
punctures in webbing; (c) unworn condition of plumage beyond disrupted area.
fore, I shall use the term thermogenic in referring to any kind of burning or
warming quahty in anting materials.)
In addition, the burning sensation produced by these freshly dug ants
seemed to vary with the season, and more of them were likely to have high
thermogenic value during summer than in winter. For instance, on February
24, 1955, when the oriole used freshly dug ants at high intensity, most of the
ants I tasted burned moderately, akhough certain others burned strongly or
222 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
not at all. But later, on May 24, when the bird anted at top intensity, the
ants were very strong in thermogenic property, so much so that a single ant
was enough to burn the whole tip of my tongue, and only an occasional one
did not burn. Furthermore, when I collected some of these ants, as the bird
applied and cast them aside, I found most of them burned strongly, although
others burned faintly or not at all.
Some of these discarded ants had been applied only once, and briefly. I
suspected the bird was applying sparingly those ants which had little or no
thermogenic property. I then collected and tasted, immediately after they
were discarded, two additional ants which, I carefully noted, had been ap-
plied several times. Each of them burned my tongue excessively. O'Rourke
(1950) discussed the variation in formic acid content of venom in several
ant species. He mentions that Stumper (1922b) showed that temperature
affected the rate of secretion in Formica ru]a and that Stumper's results gave
a QO of 2.16--that is, a rise of 10 ø C. increased the rate of secretion 2.16
times, or roughly doubled it.
Deterioration in the qualities of frozen ants was noted (p. 220) after sev-
eral weeks of storage. On November 22, 1954, I could detect no change in
ants stored since November 4; but by January 31, 1955, when the oriole re-
fused frozen ants for the first time, ! found them to be very much less sweet
and odorous and to have no burning quality whatever. The changes prob-
ably had developed earlier, because, when the bird was given a small quantity
of them on January 18, 1955, it ate all of them but applied only three. Of
even more significance, it treated only the wing tips, once possibly touching
the edge of the undertail coverts in doing so, and the right crural tract--
this last a plumage area not usually anointed.
! did not test the thermogenic property of heat-killed ants at the time of
those experiments. However, it seems probable that the heat had been suf-
ficient to lessen or destroy that property, thereby causing either poor anting
response with slow action or outright rejection, as already described. Later,
on October 20, 1955, when I did test the effects of heat, applied in the same
way, on Tapinoma sessile, I found the thermogenic value was very much af-
fected. Of 12 live Tapinoma, nine burned the tongue mildly to strongly, and
three produced no burn whatever; whereas, of 12 heat-killed specimens from
the same colony, nine gave no burn, while three burned slightly.
Unacceptable ant species differed markedly from acceptable ones in these
respects. Neither the Crematogaster, the Pogonomyrmex, nor either of the
Pheidole forms had appreciable odor, even when crushed. These species were
all rather tasteless, not at all sweet, and they did not burn.
These variables within the ant colony seemed to explain why the oriole
would eat one ant, apply the next, and fling a third away unused. I could
not be sure, of course, whether the bird's reaction to a given individual was
Lovien. ANTING IN BIRDS 223
Whitaker
determined by that ant's sweetness, odor, heating quality, or by factors yet
unknown. Evidence that the thermogenic property stimulated anting in the
Orchard Oriole may be summarized thus:
1. All ant species accepted produce a burning sensation to the human
tongue.
2. The four unacceptable ant forms were found to lack thermogenic prop-
erty.
3. Larvae and pupae of acceptable species, lacking heating property, were
rarely applied.
4. Ants of acceptable species, when subjected to heat that tended to de-
stroy the thermogenic property, brought either very low or negative anting
response.
5. Ants of acceptable species, when frozen, remained acceptable until pro-
longed storage had destroyed their thermogenic quality; once this change
occurred, frozen ants were no longer accepted.
Since the general level of thermogenic property in the ant population ap-
pears to be lower in winter, it may very well be that the oriole's tendency
toward somewhat milder antlug response in winter was due to the condition
of the ants themselves at that season. Ivor (1943) found the interest in ant-
ing pronouncedly lower among his birds in winter.
My fragmentary explorations into the properties of ants that appear to in-
duce antlug show the need for full study along this line, if we are to have
thorough understanding of anting behavior. The oriole's selectivity also
demonstrates that the behavior of birds in the presence of ants cannot be
evaluated without determination of the ant species and variability within the
colony. This is illustrated further by the instance of birds (several known
antlug species) congregating to feed on unidentified winged ants but not
performing anting actions (Worth, 1938). In this connection it is signifi-
cant that anting has not been recorded among birds following the purblind,
noisome army ants (Eciton) of vestigial sting (Schneirla, 1956; Skutch, 1954:
24; Johnson, 1954; Sutton, 1951); and that Elliott (1950), describing re-
lations of birds with red driver-ants, "probably... Dorylus (anomma)
nigricans," did not mention antlug.
Since birds are known to ant with beetles and bugs, anyone looking for
evidence of anting among birds attending swarm-raids of army ants should
bear in mind the possible role of the routed insects as antlug materials. My
Slate-colored Solitaire (Myadestes unicolor), for example, anoints itself
with certain thermogenic beetles but not with some other beetles lacking that
value, and thus far has rejected all ant species offered it.
RELATION OF FEEDING TO ANTING
As I watched the oriole perform day after day, it became clear to me that
this bird was not primarily interested in ants as food. For one thing, I soon
224 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
noted that usually more ants were discarded, with or without application,
than were eaten; and in some sessions few, if any, ants seemed to be eaten.
For instance, on June 30, 1954, during 30 minutes of top intensity anting,
the bird ate relatively few ants and I noted very few discarded without use.
Yet afterward I counted 88 damaged ants on the floor before giving up the
effort to make a full count. These, or most of them, were ants that had been
applied. When ants were offered daily, the bird's anting interest tended to
decline; and on some of those days, I did not see it eat any ants, although
it continued to ant, however mildly.
On the other hand, when the bird's interest rose, following days of low
intensity anting, I noted no corresponding increase in its intake of ants. On
one such day the bird anted eagerly; but I could be certain of seeing it eat
only three or four ants. Yet, while anting was in progress, I counted 56 in-
capacitated ants on the floor. Notes, made during periods when the bird was
given ants on many successive days, contain several remarks that the bird's
anting appetite was better sustained than its eating appetite.
No amount of anting seemed to allay the bird's hunger. When the oriole
had access to its regular food, it sometimes ate during the anting session, and
at times drank honey-water. After protracted anting sessions it commonly
fed voraciously. When ants (probably Dorymyrmex pyramicus) sometimes
were gathered with small grasshoppers in the collecting net, the oriole would
apply them before turning its attention to the grasshoppers, which it relished.
Many anting records do not show whether birds were eating ants. Most
of those that do note consumption of ants concern birds that performed ac-
tive anting. Ivor (1941; 1943) said that his birds ate a majority of the ants
they used. Poulsen (1955) stated that each ant was eaten, but that "the
birds did not always make the anting movements before eating the ant"; and
he later (1956) listed 19 species that were seen anting and sometimes dis-
carding the ants. Goodwin (1952a; 1955b) said that some birds discard all
worker ants after using them, and he described three species that habitually
went through the motions of active anting without even picking up the ants.
He (1955a) considered anting to be a "very distinct behaviour pattern, un-
connected with feeding."
Curiously, Wackernagel's Carrion Crow, which apparently did not eat ants
during the anting sessions, in one instance came back the next day to the
anting ground and ate the dead ones that remained. Huth (1951) saw a
Chaffinch (Fringilla coelebs) apply and then discard the ants. Stegmann
(letter, April 10, 1956) observed that a Brown Dipper (Cinclus pallasi) in
the Tien Shan Mountains, Siberia, in July, 1949, discarded the large brown
ants. "Holding an ant in its bill, the bird would pass it over the feathers of
the wings and of the sides of the body below the wings. This was performed
with three ants, one after the other. The ants after being used were not eaten
Lovie . ANTING IN BIRDS 225
Whitaker
but dropped anywhere." Chisholm (1944) saw no evidence of eating among
a group of Common Starlings anting in the wild. He and others comment
upon the numbers of incapacitated ants left on the anting grounds (see Brack-
hill, 1948; Thomas, 1941; Pillai, 1941; Wheeler, 1951).
It appears that birds performing passive anting usually do not eat ants
during the act. This was true of at least five of the eight Common Crows and
of the two Northwestern Crows (Corvus caurinus) for which I have been able
to find records (Frazar, 1876; Weber, 1935, and letter, May 5, 1954; Ivor,
1951; John A. Johnson, letter, September 17, 1954; Frank L. Beebe, letter,
March 15, 1954; and G. Douglas Morris, letter, November 25, 1956). Ex-
cepting one, these birds were all tame, most of them free-flying individuals.
. _ - . ,,&
.,.- .,..- , .
FIC. 4. European Jay (Garrulus glandarius) in special passive anting posture. This
species allows ants to invade plumage; it does not apply them, yet will make motions o[
picking up and applying ants to wings. Photographed by Dr. Hans LShrl, Ludwigsburg,
Germany.
The exception was the "gray-white" albinistic Common Crow which Morris
(loc. cit.) saw anting beside a trail where many large black ants were crawl-
ing around a piece of rotten wood.
An investigation of the side effects of anting and ant consumption by birds
would seem worthwhile. I have no explanation for the fact that my Orchard
Oriole occasionally held a wad of ants in its bill for quite some time, but the
behavior reminds one of the somewhat similar human use of ants as smell-
226 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
ing salts (Butler, 1897; Lhrl, 1956). The ant Oecophylla smaragdina, used
for anting, has been a regular food item among certain peoples in India, who,
it is said, use it "against fatigue and the sun's heat" (E. H. A., 1889; Long,
1901). According to Dunglison (1846:334), Formica sp. "were formerly
extolled as aphrodisiacs" and the chrysalides "are said to be diuretic and
carminative .... "Gorsuch (1934) mentioned a quail that tried to save
her nest from unidentified invading ants by eating ants "as fast as they ap-
proached until she died, possibly from formic acid poisoning."
PREENING AND BATHING FOLLOWING ANTING
The relationship of preening and bathing to anting remains obscure. The
present meager literature on this aspect of anting suggests that preening
and/or bathing after anting may prove to be pronounced only in birds that
allow ants to crawl over them and in those birds that apply vegetable sub-
stitutes with the bill.
Simmons (1955), apparently making no distinction between types of ant-
ing, has generalized: "Normal preening and bathing are almost invariable
sequels to anting." He sees in this the implication that antirig functions in
some way as a "superior preening method." However, Ivor (194.3, and letter,
April 7, 1956) has found both preening and bathing rare in his many species
anting actively with ants. Wackernagel, LiShrl (1956) and Goodwin (1947;
1951) reported bathing following antirig in certain species that permitted
ants in their plumage.
Govan (1954) noted that her Rose-breasted Grosbeak (Pheucticus Iudovici-
anus) bathed twice not very long after having anted in a skillet of sliced
onions. And she described unusual bathing by a free-living Catbird that
allowed ants to stream over its plumage during active anting. The bird in-
dulged in five brief, successive periods of anting, between each of which it
flew to a bath, where it crouched and turned about in the water, stroking
its primaries with the bill and shedding ants into the water.
Wright (1909:340) observed a case of apparent anting soon alter the bath,
in a Blue-winged Warbler (l/ermivora pinus) that dusted in a "black ant hill."
(See Dater, 1953, for details on anting in this species.) The report on three
Cardinals anting for about two and one-half hours and appearing wet at the
end of that time seems to imply that the wetness was due to the ants (Snyder,
1941). The Shackletons (1947) described two anting Indigo Buntings whose
tails and wings appeared wet, "as if drenched with liquid"; but they made
no deductions. Perhaps these, too, were cases of birds artting after bathing,
since there is nothing in these reports precluding the possibility. Here a
note by Staebler (1942) seems significant. He and George M. Sutton, notic-
ing the wet, disarranged plumage of an American Robin using Formica ex-
sectoides, believed the bird was wet from bath or the heavy dew.
Lovie. ANTING IN BIRDS 227
Whitaker
My Orchard Oriole, normally enjoying one or more baths a day, usually
had access to the bath during or immediately after anting; but its post-
anting baths were so irregular I did not keep full records. ! did, however,
make notes on 18 days when bath water was available. On seven days the
bird bathed at once after anting, in one case first dozing about two minutes
with head in scapulars. Excepting one day, these were sessions with live ants,
and anting intensities were of top (two days), high (four days), low (one
day) levels. The exceptional session was with drowned, air-dried ants, and
anting intensity was high.
On five days the oriole waited from three to 15 minutes before bathing,
once not bathing until it saw another bird do so. These performances, all
with live ants, varied from top (three days), through high (one day), to
moderate (one day) intensity.
On still another day, with live ants at high intensity, the oriole waited 30
minutes before bathing. On the remaining five days, the bird did not bathe
during the ensuing hour, and once it sunned and preened at length instead.
With one exception, these sessions were all with live ants and intensities were
high (two days), moderate (two days), low (one day). Here the exception
was with frozen ants, used at moderate intensity.
After my oriole's long and extremely exciting first experience with ants, it
flew to a perch, fluttered and shook out its plumage, preened thoroughly, and
then bathed. But ! never again noted this exact procedure. In general, it
seemed to me that bathing in this bird was too erratic to be attributed solely
to the effect of ant substances on its plumage.
Although most anting records do not mention it, preening often has been
reported in association with anting (Robien, in Stresemann, 1935b; Brack-
bill, 1948; Groff and Brackbill, 1946; Hill, 1946; Moltoni, 1948, and others).
Yet, it must be pointed out, many of these instances involved use of substi-
tutes or the invasion of the plumage by ants. Dr. Fluck wrote me that his
Blue Jays, using substitutes, preened after anting, as was true of John A.
Johnson's (MS) Common Crow following passive anting.
Aside from the preening that followed post-anting baths, preening in the
oriole was highly irregular, both in occurrence and degree. Sometimes the
bird preened not long after anting, often not at all or only much later. Oc-
casionally it shook itself briefly, scratched its head or preened a little dur-
ing breaks in protracted anting sessions. The bird did not concentrate at-
tention on the plumage parts anointed and, except when preening was brief,
it dressed the plumage as it normally would. It seemed to me that heat,
fatigue and plumage displacement were certainly as important as the local-
ized ant-ointment in motivating both bathing and preening in the oriole. I
observed that handling of the bird, which caused exertion and roughened
plumage, almost invariably induced preening, and oftimes rather prompt
228 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
bathing. Andrew (1956), studying toilet behavior of buntings (Emberiza
spp.), stated: "Disarrangement of the body feathers often seems to provoke
preening."
It appears that observers are overlooking the mechanical effects of anting.
Close observation of my oriole showed that sustained anting resulted in di-
sheveled plumage. Tripping and standing upon the tail often caused damage
to rectrices. The dabbing action of the vibrating bill regularly roughened,
and often split, webs at tips of the primaries. Damage to webbing, while
not always conspicuous, was readily discernible. Figure la shows split web
of a rectrix. Figure 3 reveals penetrations in the layered webs of the four
folded outer primaries, and particles of ant-nest earth, transferred by the
bill, adhering to margins of these openings in the web. Contour feathers
never treated during anting sometimes were disarranged by the bird's falling
and tumbling, as seen in Figure 2a. Extreme displacement of a greater wing-
covert is shown in Figure ld. The bird was photographed when in perfect
plumage, following fall molt. Several published photographs of anting birds
show split webs or the abrasive action of quill feathers pressing against the
ground (Corby, 1950; Ivor, 1956; LiShrl, 1956; Poulsen, 1956). Yet these
occurrences, as results of anting, have not been commented upon.
EXPRESSION OF THE EYES DURING ANTING
These non-spraying ants seemed to have had no effect whatever on the
oriole's eyes. I found no unusual blinking, spreading of nictitating mem-
brane, or peculiar stare in the oriole during anting, such as some observers
report. The eyes seemed to close at the instant the ant was touched to the
plumage, just as they close for deep preening. Photographs, taken a fraction
of the second before (or after?) application and while the ant is almost but
not quite touching the plumage, show the eyes open (Fig. 2c). Photographs
taken when the ant was in contact with the plumage show that the eyes are
closed or covered by the membrane (Fig. lb and Frontispiece). Corby's
(1950) photographs show two birds with eyes closed; one of them, with the
ant held some distance from the plumage, suggests the ants were a spraying
species.
I wonder whether the Mistle Thrush (Turdus viscivorus), opening and clos-
ing its eyelids (palpebre) while ants crawled in its plumage (Moltoni, 1948),
and the Baltimore Oriole, photographed in anting posture with nictitating
membrane filming the eyes (Ivor, 1956:113), may have been reacting to
ant spray in their eyes, rather than expressing enjoyment or apparent rap-
ture, as the observers interpreted. The ant (Oecophylla smaragdina), used
by birds for anting, is said to cause smarting in the eyes of jungle people
who crush masses of these insects for food (E. H. A., 1889).
When Goodwin's (1951) European Jays got acid of Formica ru/a in their
Lovle M. ANTING IN BIRDS 229
Whitaker
eyes, they would close them instantly and hop away, to stand a moment as
if pained. Various small birds, using Formica sanguinea, Lasius niger or
Camponotus pennsylvanicus (all spraying species) partly closed their eyes
just before application; but a tame Common Crow, while sitting among ants,
"closed her eyes and remained quiet for minutes at a time" (Ivor, 1951). A
tame Carrion Crow, applying and sitting among Lasius niger and Formica
ru/a, frequently drew the membrane over its eyes (Wackernagel, 1951).
LShrl (1956) said that Carrion Crows "often close their eyes with pleasure
when they are lying in an ant hill." But Liihmann (1951) believed his four
Carrion Crows closed their eyes to avoid ant spray. Poulsen's (1955, 1956)
birds, of various species, applying these last two ant species, sometimes
winked the nictitating membrane or closed their eyes, as was the case when
they were sprayed about the head experimentally with certain acid solutions
that induced anting movements. Sometimes they shook their heads or rubbed
their eyes against their shoulders.
Simmons reported that his Peking Robins (Leiothrix lutea) shook their
heads rapidly and leaped away when ant acid got into their eyes; and that
in the Magpie (Pica pica) the membrane flicked across the eyes during
antlug as a protection against acid. But Govan stated that her Rose-breasted
Grosbeak had a "trance-like stare" while using weevils found in oatmeal;
and that another such bird, applying sliced onions, held the eyes "wide and
expressionless," although afterward its eyes were misty and "almost shut."
ATTITUDES DURING ANTING
Forcefulness of the anting impulse and the apparent satisfaction derived
from the act, whether active or passive, with ants or substitutes, are men-
tioned by numerous writers. The bird's attitude has been variously described
as demonstrating enjoyment, excitement, purposefulness, or even apparent
ecstasy and intoxication. An element of compulsion seems evident in the
many cases where anting birds have ignored threat or refused to be dis-
tracted (Davis, 1944; Ivor, 1941, 1956; Groskin, 1950; Bourke, 1941b;
Lewington, 1944; and others).
The strength of the anting drive also is seen in "displacement" anting and
"anticipatory" anting (Armstrong, 1947:120; Goodwin, 1952a; Burton,
1955b, 1955c; Simmons, 1955; Poulsen, 1955, 1956; Laskey, 1949). Ac-
cording to Moynihan's (1955) definition, some of these anting acts may have
been "redirection" activities.
In its first few encounters with ants, my oriole was tense and excited. Dur-
ing its initial contact, the bird erected its crown feathers, pulled up its body
with feathers appressed, and held its tail high, as it stood among the swarm-
ing ants or ran rapidly about on the floor, snatching and applying them.
Later on, there was an eager, deliberate tenacity in the bird's manner and
230 THE WILSON BULLETIN Septelnber 1957
Vol. 69, No. 3
TABLE 3
A LIST OF BIRD SPECIES REPORTED TO ANT
Nomenclature is revised to conform with current usage. Arrangement of Passerine families and
subfamilies follows Mayr and Greenway (1956). The list includes species using ants or substi-
tutes, but not those using smoke alone. No attempt has been made to evaluate published
records.
Species Source
PItASIANIDAE
Capercaillie ( Tetrao urogallus)
Black Grouse (Lyrurus tetrix)
Ruffed Grouse (Bonasa umbellus)
Scaled Quail ( Callipepla squamata)
Chukar G41ectoris graeca)
Domestic Fowl (Gallus gallus)
Ring-necked Pheasant ( Phasianus colchicus )
MELEAGRIDIDAE
Wild Turkey (Meleagris gallopavo)
PSITTACIDAE
Unidentified parrot
Cockatoo (Kakatoe sanguinea X roseicapilla)
Turquoise Parakeet (Neophema pulchella)
STRIGIDAE
Horned Owl (Bubo virginlanus)
PIC1DAE
Wryneck (Jynx torquilla)
Flicker ( Colapres auratus)
Green Woodpecker (Picus viridis)
Golden-fronted Woodpecker (Centurus
auri/rons)
DENDROCOLAPTIDAE
Barred Woodhewer ( Dendrocolaptes certhia)
TYRANNIDAE
Ochre-bellied Flycatcher (Pipromorpha
oleaginea)
MOTACILLIDAE
Tree Pipit G4nthus trivialis)
1RENIDAE
Golden-fronted Leafbird ( Chloropsis auri/rons )
Jerdon's Chloropsis (Chloropsis /erdoni)
Chloropsis sp.
LANIIDAE
Bull-headed Shrike (Lanius bucephalus)
BOMBYCILLIDAE
Cedar Waxwing (Bornbycilla cedrorurn )
CINCLIDAE
European Dipper ( Cinclus cinclus)
Brown Dipper ( Cinclus pal[asi)
Cinclus sp.
MIMIDAE
Common Mockingbird (Mirnus po[yglottos)
Common Catbird (Durnetella carollnensis)
Reymond, 1948
Reymond, 1948
Bump, et al., 1947:272
Thomas, 1957
Reymond, 1948
Chisholm, 1944
Teale, 1953:269; MS, quoting Dash-
uta
Audubon, 1831:7; 1842:48; Sharp,
1914:65; McAtee, 1947, citing
Sharp; Allen, 1946, citing Audu-
bon
Lewington, 1944; Chishohn, 1944
Glauert, 1947
Chisholm, 1948:163-175
Mowat, 1957
Stone, 1954
F. M. Packard, MS
Allsop, 1949; Stanford, 1949
Alvarez del Toro, MS
Skutch, 1948
Skutch, 1948
Paulsen, 1956
Paulsen, 1956
All, 1936
Ringleben, in Stresemann, 1935b
Kuroda, 1947, (of related behavior
with meat)
Ivor, 1941, 1943, 1956
Creutz, 1952
B. Stegmann, MS
Heinroth, 1911a; Heinroth and Hein-
roth, 1924-1932:32; Braun, 1924;
in Gengler, 1925
C. Hagar, MS; Levon Lee, MS
Ivor, 1941, 1943, 1956; Thomas,
1946; Brackbill, 1948; Groskin,
1950; Brown, 1953; Govan, 1954
Lovie M.
Whitaker
ANTING IN BIRDS
231
TABLE 3 (Continued)
MUSCICAPIDAE
TURDINAE
Magpie-Robin ( Copsychus saularis )
Shama Thrush (Copsychus malabaricus)
Slate-colored Solitaire (Myadestes unicolor)
Veery (Catharus ]uscescens)
Gray-cheeked Thrush ( Catharus minimus)
Olive-backed Thrush ( Catharus ustulatus)
Hermit Thrush (Catharus guttatus)
Wood Thrush (Hylocichla mustelina)
Ring Ouzel ( Turdus torquatus )
European Blackbird ( Turdus merula)
Redwing ( Turdus musicus)
Song Thrush ( Turdus philomelos)
(formerly T. ericetorum )
Mistle Thrush ( Turdus viscivorus)
Clay-colored Robin ( Turdus grayi)
American Robin ( Turdus migratorius)
TIMALIINAE
Yellow-billed Scimitar-Babbler (Pomatorhinus
schisticeps )
Rusty-cheeked Scimitar-Babbler
( Pomatorhinus erythrogenys )
White-throated Laughing-Thrush (Garrulax
albogularis )
White-crested Laughing-Thrush (Garrulax
leucolophus)
Black-throated Laughing-Thrush (Garrulax
chinensis )
Gray-sided Laughing-Thrush (Garrulax
caerulatus)
Rufous-necked Laughing-Thrush (Garrulax
ruficollis )
Red-headed Laughing-Thrush (Garrulax
erythrocephalus)
Garrulax sp.
Silver-eared Leiothrix (Lelothrix argentauris)
Peking Robin (Leiothrix lutea)
Poulsen, 1956
Poulsen, 1956
Whitaker, this study
Ivor, 1941, 1943
Ivor, in Lane, 1951:177
Ivor, in Lane, 1951:177
Ivor, 1941, 1943
Ivor, 1941, 1943; Groskin, 1949,
1950; Corby, 1950, and in Hux-
ley, 1954
Reymond, 1948
Carpenter, 1945; Chisholm, 1944;
1948:163-175; Williams, 1947;
Ivor, in Lane, 1951:175; Home,
1954; Tenison, 1954; Callegari,
1955
Troschiitz, 1931, in Stresemann,
1935b; Ringleben, in Stresemann,
1935b; Poulsen, 1956
Bates, 1937; Chisholm, 1944; Gough,
1947; Wells, 1951; Fitter and
Richardson, 1951; Kent, 1952;
Poulsen, 1956
Abma, 1951; Moltoni, 1948
Alvarez del Toro, MS
Ivor, 1941, 1943, 1951, 1956; Staeb-
ler, 1942; Nichols, 1943; Van
Tyne, 1943; Davis, 1944; Law-
rence, 1945; Brackbill, 1948;
Groskin, 1950; Corby, 1950, and
in Huxley, 1954; Teale, 1953:158;
Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
Callegari, 1955
Poulsen, 1956
Callegari, 1955; Poulsen, 1956
Osmaston, 1909, 1936
Poulsen, 1956
Osmaston, 1909, 1936
Kleinschmidt, in Stresemann, 1935b;
Butler, 1910
Poulsen, 1956; Goodwin, 1955b;
Callegari, 1955
Troschiitz, 1931, in Stresemann,
1935b; Adlersparre, 1936; Steini-
ger, 1937; Ivor, 1941, 1943, in
Lane, 1951:175; Simmons, 1955;
232
THE WILSON BULLETIN
September 1957
Vol. 69, No. 3
TABLE 3 Continued)
Leiothrix sp.
Black-headed Sibia (Heterophasia capistrata)
Blue-winged Siva (Minla cyanouroptera)
Yuhina ( Yuhina nigrimentum)
Yuhina sp.
PARADOXORN1THINAE
Black-throated Paradoxornis (Paradoxornis
gularis )
SYLVIINAE
Golden-crowned Kinglet (Regulus satrapa)
MUSClCA?INAE
White-bellied Cyornis ( Cyornis tickell&e)
Rufous-bellied Niltara ( Nihava sunclara)
PACHYCEpHALINAE
Rufous Whistler ( Pachy cephala rujiventris )
Little Shrike-Thrush ( Colluricincla parvula)
ZOSTEROPIDAE
Western White-Eye (Zosterops palpebrosa)
MELIPHAGIDAE
Yellow-eared Honeyeater (Meliphaga lewini)
EMBERIZIDAE
EMBERIZINAE
Brazilian Cardinal (Paroaria capitata)
Black Seedeater ( Sporophi[a aurita )
Red-eyed Towbee (Pipi[o erythrophtha[rnus)
Brown Towbee (Pipi[o/ascas)
Slate-colored Janco (Junco hyema[is)
Harris Sparrow ( Zonotrichia querula )
White-crowned Sparrow (Zonotrichia
[eucophrys)
White-throated Sparrow (Zonotrichia
albicollis )
Fox Sparrow (Passerella iliaca)
Song Sparrow ( Melospiza melodia)
CARDINALINAE
Common Cardinal ( Cardinalis cardinalis)
Rose-breasted Grosbeak (Pheucticus
ludovicianus )
Black-headed Grosbeak (Pheucticus
melanocephalus )
(Pheucticus rnelanocephalus X P.
ludovicianus )
Buff-throated Saltator (Sahator maximus)
Indigo Bunting (Passerina cyanea)
Lazuli Bunting ( Passerina amoena)
Poulsen, 1956; Goodwin, 1955b;
Callegari, 1955
Kleinschmidt, in Stresemann, 1935b;
Lorenz, in Stresemann, 1936
Troschiitz, in Stresemann, 1935b;
Poulsen, 1956
Poulsen, 1956
Osmaston, 1909, 1936
Poulsen, 1956
Poulsen, 1956
Davis, 1939
Poulsen, 1956
Poulsen, 1956
Bourke, 1941a, 1941b
Sedgwick, 1946
Poulsen, 1956
Bourke, 1941a
Poulsen, 1956
Skutch, 1948, 1954
Van Tyne, 1943; McAtee, 1944;
Corby, 1950
Paroni, 1954, (of an attempt)
Ivor, 1941, 1943; Bagg, 1952
Ivor, 1943
Ivor, 1941, 1943; Paroni, 1954
Ivor, 1941, 1943; Teale, MS
Ivor, 1941, 1943
Nice and Ter Pelkwyk, 1940; Ivor,
1941, 1943; Nice, 1943; Mayr,
1948; Groskin, 1950
Edwards, 1932; Ivor, 1941, 1943,
1956; Snyder, 1941; Sprunt and
Chamberlain, 1949:510-511;
Chamberlain, 1954, quoting Mc-
Atee
Ivor, 1941, 1943, 1956; Govan, 1954
Ivor, 1943
Ivor, MS
Skutch, 1948, 1954
Ivor, 1941, 1943; Shackleton and
Shackleton, 1947; Poulsen, 1956
Poulsen, 1956
Lovie M.
Whitaker
ANTING IN BIRDS
233
TABLE 3 Continued)
Poulsen, 1956
Orange-breasted Bunting (Passerina
leclancherii )
TANAGRINAE
Superb Tanager ( Calospiza jastuosa)
Blue-breasted Tanager (Calospiza
cyanoventris )
Blue-necked Tanager ( Calospiza cyanicollis)
Scarlet Tanager ( Piranga olivacea )
Summer Tanager (Piranga rubra)
Red-throated Ant-Tanager ( Habia gutturalis )
COEREBINAE
Blue Sugarbird (Dacnis cayana)
PARULIDAE
Blue-winged Warbler ( Vermivora pinus)
ICTERIDAE
Brown-headed Cowbird ( Molothrus ater)
Boat-tailed Grackle ( Cassidix mexicanus )
Common Grackle ( Quiscalus quiscula)
(includes Q. q. stonei, Q. q. aeneus,
Q. versicolor)
Tinkling Grackle (Quiscalus niger)
Baltimore Oriole (Icterus galbula)
Orchard Oriole (lcterus spudus)
Troupial ( lcterus jamacaii)
lcterus sp.
Black-throated Oriole (lcterus gularis)
Streak-backed Oriole (Icterus pustulatus)
Red-winged Blackbird ( zl gelaius phoeniceus )
Common Meadowlark ( Sturnella magna)
Bobolink ( Dolichonyx oryzivorus)
FRINGILLIDAE
FRINGILLINAE
Hawfinch ( Coccothraustes coccothraustes)
Chaffinch ( Fringilla coelebs )
Brambling ( Fringilla montijringilla )
CARDUELINAE
Evening Grosbeak ( Hesperiphona vespertina)
ESTRILDIDAE
Red-browed Waxbill ( Estrilda temporalis )
PLOCEIDAE
Bubalornis ( Bubalornis alblrostris )
House Sparrow (Passer domesticus)
Chestnut Weaver (Ploceus rubiginosus)
Yellow-shouldered Widow-Bird (Coliuspasser
liZaCFOCeFC1ZS)
Whydah (Coliuspasser ardens)
Long-tailed Widow-Bird ( Diatropura progne )
Jackson's Widow-Bird (Drepanoplectes
jacksoni)
Bengalese Finch (Munia striata), domestic
form
Poulsen, 1956
Sick, 1957
Sick, 1957
Groskin, 1943, 1950
Thomas, 1941
Alvarez del Toro, MS
Poulsen, 1956
Wright, 1909; Dater, 1953
Nice, 1945; Hebard, 1949
W. W. Worthington, MS
Ivor, 1941, 1956; Parks, 1945; Rob-
inson, 1945; Groff and Brackbill,
1946; Hill, 1946; Brackbill, 1948;
Laskey, 1948; Nice, 1952; Teale,
1953:158, 170; Poulsen, 1956
Gosse, 1847:225
Ivor, 1941, 1943, 1956
Whitaker, this study; Ivor, MS
Poulsen, 1956
Goodwin, 1953; Simmons, 1955
Alvarez del Toro, MS
Alvarez del Toro, MS
Nero, 1951; Teale, 1953:158; Poul-
sen, 1956
Ivor, MS
Ivor, 1943; Nice, 1943
Poulsen, 1956
Longhurst, 1949; Goodwin, 1951,
1955b; Huth, 1951; Poulsen, 1956
Poulsen, 1956
Ivor, 1941, 1956
Givens, 1945, (of smoke-bathing and
attempted use of ants)
Poulsen, 1956
Davis, 1945; Wheeler, 1951; Com-
mon, 1956
Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
Ivor, MS
234
THE WILSON BULLETIN
September 1957
Vol. 69, No. 3
TABLE 3 (Continued)
Taha Bishop (Euplectes taha)
Orange Bishop (Euplectes ]ranciscana)
Fire-crowned Bishop (Euplectes hordeaceus)
STURNIDAE
Long-tailed Glossy Starling (Lamprotornis
caudatus )
Glossy Starling ( Lamprotornis chalybaeus )
Superb Starling (Spreo superbus)
Rose-coloured Starling ( Sturnus roseus )
Common Starling ( Sturnus vulgaris )
Pied Starling (Sturnus contra)
Indian Mynah ( Acridotheres tristis )
Chinese Jungle Mynah (Acridotheres
cristatellus )
Indian Jungle Mynah (Acridotheres ]uscus)
Bank Mynah ( Acridotheres ginginianus)
Indian Grackle ( Gracula religiosa)
DICRURIDAE
Drongo Dicrurus sp.
GRALLINIDAE
Magpie-Lark ( Grallina cyanoleuca)
Apostle-Bird ( Struthidea cinerea)
CRACTICIDAE
Australian Magpie ( Gymnorhina dorsalis )
PARADISAEIDAE
Green Catbird ( Ailuroedus crasslrostris)
Satin Bowerbird ( Ptilonorhynchus violaceus )
CORVIDAE
European Jay ( Garrulus glandarius )
Lanceolated Jay ( Garrulus lanceolatus)
Adlersparre, 1936
Adlersparre, 1936; Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
OLD WORLD:
Hoyningen-Huene, 1869; Heinroth,
1911a; Floericke, 1911; Gengler,
1925; Hampe, in Stresemann,
1935b; Moncrieff, 1935; Scheid-
ler, in Stresemann, 1936; Chis-
holm, 1944, 1948; Gregory, 1946;
Baggaley, 1946; Tebbutt, 1946;
Hobby, 1946; Armstrong, 1947:
120; Prideaux, 1947, (of using
smoke); Williams, 1947, 1948;
White, 1948, (of using smoke);
Abma, 1951; Wheeler, 1951; Fit-
ter and Richardson, 1951; Good-
win, 1951, 1955a, 1955b; IJzen-
doorn, 1952a, 1952b; Moltoni,
1952, quoting Binelli; Simmons,
1955; Poulsen, 1955, 1956
NEW WORLD:
McAtee, 1938, quoting Kalmbach;
Pearson, 1938; Brackbill, 1948;
Ivor, in Lane, 1951:175; 1956;
Teale, 1953:158, 159, 199
Poulsen, 1956
Chisholm, 1935a, 1944, 1948:163-
175; Pillai, 1941; Wheeler, 1951
Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
Poulsen, 1956
Fletcher, 1937
Chisholm, 1944; Galloway, 1948
Chisholm, 1944, 1948:163-175
Sedgwick, 1947
Poulsen, 1956
Chisholm, 1944
Robien, in Stresemann, 1935b;
Hampe, in Stresemann, 1935b;
Goodwin, 1947, 1951, 1952a, 1953a,
1953b, 1955b; Li3hrl, 1952, 1956;
Simmons, 1955; Burton, 1955c;
Poulsen, 1955, 1956
Goodwin, 1952a, 1953a, 1953b, 1955b
Lovie M.
Whitaker
ANTING IN BIRDS
235
TABLZ 3 (Continued)
Lidth Jay ( Garrulus lidthi)
Blue Jay ( Cyanocitta cristata)
Beechey's Jay ( Cissolopha beecheli)
Green Magpie ( Kitta chinensis
Red-billed Blue Magpie (Kitta
erythrorhyncha)
Azure-winged Magpie (Cyanopica
cyanus)
Magpie (Pica pica)
Tree-Pie ( Crypsirina bayleyi)
Rook (Corvus [rugilegus )
Common Crow (Corvus brachyrhynchos )
Northwestern Crow (Corvus caurinus )
Carrion Crow (Corvus corone)
(Hooded Crow, C. corone cornix, included)
Raven (Corvus corax)
Kuroda, 1947, (of related behavior
with acorn: conpare with Good-
win, 1952a)
Baskett, 1899:243; Ellicott, 1908;
Ivor, 1941, 1943, 1946, 1956; Lane,
1943; Buell, 1945; Fluck, 1948 and
MS; Laskey, 1949; Corby, 1950;
Davis, 1950; Miller, 1952; Anon.,
1952; Teale, 1953:158, 168; Nice,
1955a; Poulsen, 1955, 1956
Goodwin, 1952a, 1953a
Goodwin, 1953a, 1955b; Poulsen,
1956
Goodwin, 1952a, 1953a, 1955b; Poul-
sen, 1956
Nonomiya, 1935; Goodwin, 1953a,
1955b
Heinroth, 1911a; Funke, 1912;
Chisholm, 1940, 1944; Reynolds,
1946; Schierer, 1952; Goodwin,
1953a, 1955b; Simmons, 1955
Osmaston, 1909, 1936
Prideaux, 1947, (of using smoke);
McMeeking, 1949, (of using
smoke); Chappell, 1949; Good-
win, 1953a, 1955b; Burton, 1955a,
1955b
Frazar, 1876; Weber, 1935; lvor, in
Lane, 1951:17,%177
F. L. Beebe, MS
Heine, 1929; Laven, 1931; Ringle-
ben, in Stresemann, 1935b;
Scherping, in Stresemann, 1936;
Condry, 1947; Coombs, 1947;
Wells, 1950; Liihmann, 1951;
Wackernagel, 1951; Goodwin,
1953a, 1955b; Lhrl, 1956
Jacobsen, 1911
little or no excitement. Distractions sufficient to prevent or interrupt sunning,
bathing or feeding rarely kept the oriole from anting. On a few occasions
it called chak, but never sang during anting sessions.
SEASONAL FLUCTUATIONS IN ANTING RESPONSE
The Orchard Oriole performed in every month of the year. In the warmer
months (March to August), when opportunities were most frequent, anting
intensity ranged from top level to very low. In the periods from September
to February it ranged from high to very low. Experiments did not test sea-
sonal differences adequately, because of variation in conditions and species
of ants offered. However, it is noteworthy that even live ants did not in-
duce top intensity anting in September through February but did so from
March through July. On the other hand, the oriole anted at high intensity
in September and in November through February. As noted earlier, seasonal
236 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
changes among the ants themselves appeared to affect intensity of anting
response.
As would be expected, a breakdown of American and Canadian records
of anting in the wild shows by far the greatest occurrence during May through
October, with the highest incidence in August. I find no reports for De-
cember to March save in captive birds, except the North Carolina record of
a Golden-crowned Kinglet (Regulus satrapa), using unknown anting ob-
jects in a tree in January (Davis, 1939). Captives appear to ant throughout
the year. But Ivor, in Canada, seems to be the only observer who has given
close attention to this feature. His birds, representing about 32 American
and two or three exotic species, exhibited but little interest in ants except
from late April through July.
It seems possible that winter anting may take place in the warmer parts
of the United States, where ants are active above ground on sunny days. Ob-
servers in these regions should watch for the behavior. But, as Herbert L.
Stoddard, Thomasville, Georgia, suggests to me, care should be taken that
feeding on weed seed in the nests of harvester ants is not mistaken for anting.
There are relatively few records of anting from southern United States for
any season, and only one conclusive record (Levon Lee, letter, May 13, 1956)
from the area between western Texas and Oakland, California, as far as I
have been able to discover. This lacuna in the Southwest, first noted by
Kelso (1949), probably is more apparent than real, since several species
breeding there are now known to ant (see Table 3). Because anting some-
times is inconspicuous (Nice, 1945), or resembles preening (Simmons, 1955,
Poulsen, 1956), it no doubt often goes unrecognized. Jean Graber has called
my attention to an unquestionable case of anting by a Cardinal reported
as food-gathering (Edwards, 1932).
Considering how very little we know about anting, it might prove help-
ful if operators of banding stations would place ant colonies in the traps.
Confined by water barrier, the captive colony requires little care and could
be a means of obtaining valuable information, particularly as to general
health and parasitism among birds anting in the natural state. Obviously
ants chosen for this purpose must be of a known acceptable species.
Some workers dismiss all observations on captive birds, apparently be-
cause of such factors as inactivity, disease or unnatural diet. Others have
been equally reluctant to accept reports of anting among free birds of groups
long believed to be non-anting species, such as the Psittacidae or Picidae,
simply because captives did not ant. These positions become untenable when
we consider the many species, first known to ant in captivity, that are now
known to ant in the same way in the wild, and vice versa. It may 'indeed be
true that the captive bird is more prone to ant than is its counterpart in na-
turelack of normal energy outlets alone might make the difference. But
Lovie M. ANTING IN BIRDS 237
Whitaker
it remains to be demonstrated that captivity per se either causes or sup-
presses antirig, or modifies characteristic antirig patterns.
Antirig in captive birds free of obvious ectoparasites has been reported by
Poulsen, Adlersparre, Heinroth (1911a), Ivor (letter, May 5, 1954), Scheid-
ler (in Stresemann, 1936), and others. Parks (1945) found no parasites on
the Common Grackle he trapped immediately after anting. Beebe, Weber,
and J. A. Johnson advised me that their several crows had no obvious ecto-
parasites, and Beebe adds that his birds had been dusted with a rotenone
preparation prior to antirig. Frazar's (1876) two Common Crows and the
two or three Australian cases involving domestic fowl and free Starlings
(Sturnus vulgaris), cited by Chisholm (1944), may be almost the only re-
ports of antirig in obviously infested birds. It is indeed interesting to see
that an Old World oriole (Oriolus melanocephala), infested with mites, did
not ant, although it ate the ants (Poulsen, 1956).
The plain fact of the matter is that we know almost nothing of the physical
condition of antirig birds. No one seems to have made skin scrapings or any
close examination for minute ectoparasites. Endoparasites as a cause of ant-
ing have received scant attention. I have been unable to find a single in-
stance of dissection study on an artting bird, yet at least two ants, one of
them Pheidole sp., are known to be intermediate hosts for two types of cysti-
cercoids in chickens (Jones and Horsfall, 1935; Eichler, 1936b). Since endo-
parasites may, through lack of intermediary hosts, be lost in the captive bird,
they should be considered in connection with the onset of negative antirig
response sometimes seen in captives.
DIscussION: THE ANTING OB.IECTIVE
Study of the antirig pattern of the Orchard Oriole revealed that during
intense artting, more often than not, the bird was dabbing ants in the region
of the vent. The undertail coverts and very bases of the rectrices certainly
were treated, if not the vent itself. As results of the bird's vigorous appli-
cations, I found that ant scent was strongest in this region and that the under-
tail coverts at times became mingled with uppertail coverts.
Tallies of applications indicated preferential treatment in that area also;
however, these counts necessarily were incomplete because of the bird's
rapid action and occasional obstruction of my view. One of many tallies
showed 46 applications to basal half or less of tail, including 15 to the under-
tail coverts alone; and 22 applications to distal one-third or less of primaries,
including a number wherein the basal part of the tail shared in the treatment,
as will be explained.
Even when the wing tip was treated, it often seemed that the oriole really
was aiming at the undertail region, and that the wing interfered. Almost
invariably, as the bird began to reach back with the ant, it simultaneously
238 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
tucked the posteror body down and under, until crissum at times touched
the heels, and rectrices were brought forward on one side of, or between, the
feet. In this latter posture, the bird sometimes would anoint the lower belly
and the undertail coverts between its heels. Regardless of the area treated, the
bird usually reached around on one side, bringing the tail around on that
same side. In doing this, the wing on that side would be folded high upon the
body, with remiges out of the way behind the tail (Fig. la). Or again, the
folded wing might be held lower, along side and flank, so that wing tip lay
against the ventral surface of the tail, as the tail was tucked under and
pressed forward. This pressure of tail upon wing was strong (Frontispiece;
Figs. ld and 2c), and often caused rectrices to interlock with remiges (Fig.
lb). Though he does not discuss it, one of LShrl's (1956) photographs of
an anting Carrion Crow demonstrates a similar interlocking of wing and
tail in a passively anting species. In much the same manner, some passively
anting birds at times will apply ants with the bill, as his bird is shown doing.
During the oriole's treatment of the wing tip, when tail was pressing against
it, basal parts of rectrices (including undertail coverts) at times received
some of the anointment, thus increasing the proportion of applications af-
fecting the undertail region. This situation is seen in the Frontispiece. The
lacerated webs shown in Figure 3, together with the extreme displacement
of undertail coverts, already described, indicate that the oriole's bill some-
times penetrated deeply into the plumage. All of this strongly suggests that,
when anting in this tail-to-wing posture, the bird sometimes thrust the ant
through the wing tip and onto the under surface of the tail. Certainly, at
such times, I regularly noticed that the dabbing applications of the vibrating
bill involved both the wing tip and the adjacent areas of the tail, including
the undertail coverts. In several photographs of actively anting birds (Corby,
1950; and in Huxley, 1954; Poulsen, 1956; Ivor, 1956) this same juxtaposi-
tion (and wing-tail anointment?) is illustrated though unemphasized. How-
ever, Poulsen (op. cit.) did state that it "often looks as if" the tail is treated
and that in some cases among starlings, weavers and babblers, he has seen
the bill movements "proceed to the tail, which is held close to the wing."
This is not to say that the oriole never anointed the wing tip when the
wing was slightly spread and held just clear of the body and tail. It did so
regularly; but instead of holding the opened wing out to the side as some
birds do, the position of the wing was like that assumed for stripping the
outer primaries during preening. Furthermore, this posture was seen less
often than either of the other two anting positions just described, and at such
times the applications themselves often were atypical. That is, the dabbing
action of the bill often was so slow or so brief as to give me the impression
that the bird was anting "absent-mindedly"--or possibly confusing anting
with preening. Frequently, when anting in this position, the bird would dab
Lovie M. ANTING IN BIRDS 239
Whitaker
the ant about on the wing tip a time or two and then stop, as if confused.
Aberrant anting action was seen again in the oriole's occasional treatment
of the crural tracts. Instead of dabbing forcefully, as it did in anointing
other areas, the bird always would play the ant about on these feathers very
lightly, briefly and, it seemed to me, ineffectually and sometimes accidentally.
Ivor (1943) has described somewhat similar action in young Wood Thrushes
(Hylocichla mustelina) that sometimes anted at breast, abdomen and flanks,
without actually touching those parts; and Brackbill (1948) noted an
American Robin that twice seemed to dab at, but not actually touch, its
breast during anting. Poulsen (1956) mentioned that an Indigo Bunting,
while applying ants, intermittently made incipient movements of picking up
and applying an ant.
These unusual motions, as well as the oriole's sometimes odd wing treat-
ment, might well betoken uncompleted anting acts, occurring when the in-
dividual ant used happened to have an inadequate amount of stimulant.
Conclusive evidence might be obtained through controlled experiments with
spraying ants which previously have been forced to substantial ejection of
their defense fluid and then washed. It seems quite possible that such de-
pleted ants could produce significant differences in a bird's anting actions.
Poulsen (1956:281) noticed that "birds anted much less with ants [spraying
species] which had been kept in a sack for some days and therefore were
less active." (Might the rate of venom secretion in the glands of these ap-
parently unfed captive ants have been lower?) My experience with the
Orchard Oriole leads me to believe that more precise experimentation with
heat-killed ants might show correlation between the level of thermogenic
property in the ant and the plumage area treated. I suspected the oriole of
using the more strongly stimulating ants on the undertail coverts and bases
of rectrices.
It must not be supposed from the foregoing discussion that the oriole treat-
ed only the proximal half of the tail. Frequent anointment of the distal por-
tion was extended at times to the very tips of rectrices. Commonly, however,
an application to the distal part of the tail began as an application to under-
tail coverts or bases of outer rectrices, the bird merely continuing to reach
and dab farther along the tail before stopping and straightening up.
The thermogenic property of ant species accepted by the oriole presumably
is due to an irritant in the secretions of their anal glands. Ants of the sub-
family Dolichoderinae, which includes all of the acceptable ant species
studied here, are known to smear their defensive secretions on enemy ants
with frequently fatal effect (Wheeler, 1910:45). It seems, however, that
myrmecologists have not determined the irritating constituent in those ant
species my bird used. The unpleasantly odorous butyric acid, which these
ants are believed to produce in quantity, is non-caustic. A free acid in butter,
240 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
it has been noted also in meat juice, perspiration and excrementa. It would
be helpful to know whether all insects used for anting produce burning sen-
sations. Ants producing formic acid in quantity no doubt do burn, since
this is a strong caustic. Many birds have used ants of this type. It appears,
therefore, that a sensation of heat is a probable factor in most anting situa-
tions, as Burton (1955c) surmised.
In speculating on causes of anting, it would be well to keep in mind these
points: (a) the fact that a bird sometimes will persist despite interruptions
and threat situations, or even continue anting to a state of obvious fatigue;
(b) that in captives, at least, the amount of anting varies among species and
among individuals of a species; (c) that sudden, unexplained, even apparently
permanent abstinence has been seen in captives; (d) that not all captives of a
species will ant; (e) that apparently not all, or even most, individuals of a
species ant in nature, at least not with any regularity; (f) that active and
passive anting are not necessarily mutually exclusive--some passively anting
species exhibit elements of active anting, and vice versa; but among birds
under 10 inches in length, passive anting elements seem to be unusual.
Gross ectoparasltes as a cause of the orlole's pronounced anting of the
undertail region would have to be ruled out. There was no evidence what-
ever of such parasitism. But it is possible that itch mites (Acarina) might
have been present in skin of the vent and its environs, especially mites in
the pockets at feather insertions. Tapeworms (Cestoda) or roundworms
(Nematoda) emerging from the vent, or flukes (Trematoda), sometimes
known to encyst beneath skin in that region, might possibly cause itching.
(For discussion of parasites see Rothschild et al., 1952:39-242; Peters, 1930,
1933, 1936; Boyd, 1951; Eichler, 1936b.) Aside from attention given this
area during anting, ! saw nothing in the oriole's behavior suggestive of ir-
ritation. It did not pick, preen unduly or rub its posterior. Yet it did thrust
ants among the bases of rectrices and, when treating the outermost few of
them, was seen to apply ants at the feather insertions. (Also, if parasites were
the sole cause of the bird's anting, then these must have been host-specific
ones, for the oriole's companion, the Painted Bunting, has never anted.)
There are many records of repeated applications to undertail coverts, "base"
or "root" of tail; and in some other cases such treatment seems implied
(Bates, 1937; Goodwin, 1953a; Groskin, 1950; Nice and Ter Pelkwyk, 1940;
Osmaston, 1909, 1936; Thomas, 1946; Staebler, 1942; Snyder, 1941; Brack-
bill, 1948; Davis, 1950; Home, 1954; Tebbutt, 1946, and others.)
The pleasure principle, on the other hand, seems a more likely basis for
theorizing on the oriole's behavior. If, as it appears, the bird's mouth parts
are sensitive to thermogenic properties in ants, it seems reasonable to sup-
pose the vent, and perhaps the skin of the undertail region, would be similar-
ly sensitive. Indeed, Simmons suggests that the area of the vent must be a
Lovie M. ANTING IN BIRDS 241
Whitaker
most sensitive spot. Ant secretions might cause a peculiarly pleasurable
sensation of warmth, possibly with an element of the masturbatory in it. Auto-
eroticism is known in domestic parakeets, and, according to Armstrong
(1947:160), also in parrots, ruffs, avocets, sage grouse and penguins. One
wonders whether the posture of Carrion Crow on its perch, after having anted
on the ground, could be a result of heat, fatigue, mild sexual stimulation, or
FIC. 5. Carrion Crow (Corvus corone) in ordinary passive anting posture typical of
certain larger coryiris. Note spread tail, somewhat pulled toward left wing, and lifted
contour plumage. Photographed by Dr. Hans Lhrl, Ludwigsburg, Germany.
some combination. Goodwin (1953a; 1955b, figure) showed the bird in
relaxed attitude, head tilted downward, tail drooped, and both wings hanging
in front of, and well below, the perch.
Might there be sexual significance in those instances of both free and
captive birds bringing the vent into close proximity to ants, either by direct
application in that region or by holding the vent near the ground? Several
of the American Robins, performing active anting, sometimes crouched,
rotated or rubbed the body or breast upon the ground, or sat as if holding
vent to ground (Nichols, 1943; Van Tyne, 1943; Brackbill, 1948). lvor
(letter, November 3, 1954) informs me that two of 12 Robins regularly
crouched but that the others never took that position when anting. Certain of
Poulsen's thrushes ( Turdus migratorius, T. musicus and T. philomelos ), while
242 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
applying ants, sometimes assumed passive anting posture. He writes that
they "suddenly fluffed their feathers and spread both tail-feathers and wing-
feathers, making 1-3 strokes with a single ant, which was then discarded. All
the while they were sitting on the ground and allowing the ants to crawl on
them without removing them .... "Lawrence (1945) cited a rather similar
case for migratorius. Comparable photographs showing American Robins in
partial crouch with this double-wing-spread appear in fvor (1956) and
Poulsen (1956). (Might these variations evident in migratorius be related
to sex difference?)
Posture that would seem to expose the area of the vent to ants also has
been described for some other species. Poulsen, as well as Brackbill, noted
Common Grackles that repeatedly sat on the ground, with tails stretched out
behind, while applying ants. A captive Rook flopped down on the anting
area and raised its tail while the ventral body was flat on the ground; and
a captive Carrion Crow pressed its tail against the earth while sprawling on
the anting area (Goodwin, 1953a). Both of these last birds, though using
passive anting posture, at times applied ants with the bill. Wackernagel's
Carrion Crow applied ants but also lay flat on its belly with wings spread.
While lying down, it several times struck here and there with its fanned tail.
Once it sat, as if brooding, and rubbed the "anal area" on the ground. Some-
times it interruped anting to go to the turf and, with widely spread tail, drag
its plumage through the grass. Simmons states that in the Magpie the ant
sometimes quite definitely is "rubbed in the area of the vent" and he suspects
this is true of many smaller birds whose quick motions are hard to observe
in detail. Osmaston's (1936) birds, three or four species, using bugs (Rhyn-
chota), rubbed them only "near the anus."
The foregoing descriptions relate to individuals that performed more or
less active anting. As for those that stand, squat or lie and make few or no
applications with the bill, the skin of the ventral body, including the vent,
would seem especially vulnerable to ants and ant spray, particularly when the
contour plumage is fluffed, as often seems the case during passive anting.
And here we should bear in mind that ant spray may carry eight inches or
more.
Captive European Jays showed "apparent attempts to bring the ventral
areas in contact with ants by dragging them along on the ground" (Goodwin,
1951). These birds customarily brought forward or depressed the tail,
spread both wings forward with convulsive, shuddering spasms, and con-
stantly ran the bill down the wing quills without actually applying ants or
even picking up ants. The insects swarmed up their legs and into their
plumage. Goodwin's (1955b) sketches of some characteristic anting postures
show this species in upright, almost penguin-like attitude, with undertail
coverts (and vent?) apparently touching the ground, tail flat on the ground
Lovie M. ANTING IN BIRDS 243
Whitaker
behind, and the posterior edges of the opened wings just brushing the ground.
Like posture in this jay is shown here (Fig. 4) and in other photographs
(LiShrl, 1956; Burton, 1955c) and descriptions (Goodwin, 1947; 1952a;
Poulsen, 1956; Robien, in Stresemann, 1935b). Incidentally, Burton (loc.
cit.) demonstrated its occurrence in response to substitute materials.
Scherping (in Stresemann, 1936) observed that a tame young Carrion
Crow lying with spread wings on an ant heap never did apply the ants.
Condry's tame young bird of this species behaved in much the same way but
drooped the head in a "swooning" position until beak touched the ground, and
sometimes it put ants on its back with the bill. Chisholm (1944) referred
to a Magpie-Lark (Grallina cyanoleuca) that sprawled on the ant mound as
if incapacitated. A young captive Hooded Crow lay with half-spread wings,
flapped them as if bathing, but did not apply the ants (Coonabs, 1947); and
a tame Mistle Thrush pressed itself against the ground, wings spread, while
ants crawled on its body (Moltoni, 1948).
Liihmann's Carrion Crows ruffled their plumage, lowered their breasts
onto the ant nests, and made bathing motions. Lest this appear to have been
mere dust-bathing, it should be added that Liihmann remarks that he was
never able to see even an attempt at anting whenever few ants were present
at the nest surface. Beebe's Northwestern Crows sprawled, almost as if dead,
while ants crawled upon them, except that the head was held up to one side,
exposing apteria at the back and shoulders.
As Lihrl and Condry state of the Carrion Crow, John A. Johnson's (letter,
September 17, 1954) Common Crow permitted ants to cover it from neck to
tail. When picked up from the ant bed before surfeit, the bird would return
to the ants at once. It would stand until the insects began crawling up its
legs, then would lie down on one side, with fluffed plumage and lifted wing,
or again, on its breast with both wings slightly spread. Sometimes it "spread
feathers at base of the tail" with its beak, apparently "to let ants reach the
skin" but it was never seen to apply ants with the bill.
The above cases of passive anting are not altogether different from the
behavior of certain mammals with ants. Bagg (1952) watched a gray squirrel
(Sciurus carolinensis) roll and tumble on unidentified ants, and occasionally
crawl on its belly across the nest. Swanson (1956) described a "timber
squirrel" that rubbed its belly and head against a spot on the trunk of a
maple tree where ants were feeding on the sap. And Chisholm (1948:163-
175) mentioned the case of a domestic cat's (Fells domesticus) ecstatic con-
tortions on a rubbish heap where ant-debris from a collector's can had been
emptied. It may be significant that a fox and a squirrel evidently did not
behave in any unusual way while eating termites (Stewart, 1888), and that
anting with termites, which apparently do not liberate defense fluids, appears
to be unknown in birds.
244 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
From cases reviewed here--and others could be citedit seems clear that,
as a result of either active or passive anting, a bird can receive ant substance
upon the skin of the undertail region in proximity to the vent and probably
on the vent. ! believe further study may show a main focus for stimulation
in many, if not all, anting birds to be the undertail region, probably the
vent, hence the peculiar positions of the tail which sometimes cause tripping
or falling.
In analyzing the literature pertaining to active anting, one is struck by
the comparative rarity of instances involving application to dorsal body areas,
or even dorsal surfaces of the quill feathers--these latter being favorite
targets ventrally, according to most observers. Furthermore, such records
usually indicate that the dorsal applications were few or that, in the main,
the bird was anointing ventral areas of the wings, tail or body. Also, one
notes that rump and upper tail coverts are mentioned more often than the
other dorsal parts. One citation of application to the back in a young Dipper
(Cinclus sp.) seems due to error in translation and should read belly (Bauch)
instead (McAtee, 1938, citing Heinroth, 1911a). Anointment of the anterior
body appears to be rare, except in grackles, and almost invariably is accom-
panied by treatment of wings and tail. It is noteworthy that three species of
grackles (Icteridae), which are prone to anoint various parts of the body,
wings and tail, usually have used one or more of eight substitute materials.
Indeed, with these birds, the use of ants would seem to be the exception
rather than the rule (see citations in Table 3). The observations of W. W.
Worthington (letter, November 24, 1956) on two species of grackles in
nature relate to use of limes and lemons on the wings, tail, breast, neck and
scapulars, and are thus similar to the reports of several other persons. Poulsen
(1956), ignoring passive and active anting as the two basic types of the be-
havior, recognized five types of anting, to one of which he assigned only his
three Common Grackles (Quiscalus quiscula), on the basis of their applying
ants to the breast, scapulars, rump and upper tail coverts, in addition to the
wings.
It is significant also that generally those birds (seldom under 10 inches)
which permit massive invasion of the plumage by ants seem not to apply
ants with the bill, or seem to do so rather casually or sparingly. Since a good
many of the commonly used spraying ants are capable of ejecting their spray
with considerable force, may not these birds obtain the desired stimulation
(of the ventral body skin) with no effort on their part other than that of
standing, squatting or lying with spread wings and sometimes raised contour
plumage? (Compare descriptions and the illustrations of Goodwin, 1952a,
1955b; Coombs, 1947; Condry, 1947; Liihmann, 1951; Wackernagle, 1951,
with those of Brackbill, 1948; Groskin, 1950; Ivor, 1941, 1943, 1956.)
In a number of cases, birds behaving this way were known to settle down
Lovie M. ANTING IN BIRDS 245
Whitaker
among the ants only after ants had crawled up their legs (and stimulated
body skin?). Wackernagel's bird went to spraying ants and stalked back
and forth (increasing the quantity of ant spray?) before settling down
among the insects or applying them with the bill. Robien's (in Stresemann,
1935b) European Jays trod upon ants as if to increase the spray; Condry's
Carrion Crow, when ants crawled up its legs to the feathers, sat down among
them like a brooding hen.
I have described the oriole's way of treating the distal portion of the tail
in a follow-through of applications first made directly to undertail coverts
and basal portions of rectrices. Could this extended dabbing, out along the
ventral side of the tail, possibly be reflexive action, due to ant substances
warming the skin of the undertail region? Might such stimulation cause
some of the special movements of wings and tail reported in certain species?
The idea of special movements being reflex consequences of thermogenic
agents on the skin may sound far-fetched unless we recall that some of these
actions evidently occur after ants have been applied or ant spray, presumably,
has reached the skin.
Goodwin (1947, 1952a, 1953a) clearly showed European Jays standing
among spraying ant species and assuming their double-wing-spread posture
only after ants had swarmed up their legs, in one case after they had reached
the ventral body plumage. He and Poulsen both found that this species, the
Green Magpie (Kitta chinensis), and the Red-billed Blue Magpie (K. erythro-
rhyncha) would advance both opened wings, accompanied by convulsive
shudderings. Each of these species let ants swarm into the plumage; none
actually applied ants, although they went through the motions of doing so.
Poulsen's Blue Sugarbird (Dacnis cayana) took like posture. He (1956:274)
wrote "This species picks up an ant [spraying species] in its bill, and very
rapidly it rises in an almost vertical position with spread tail and moves both
wings forward so that they touch each other while quivering, and the head
is moved downwards among the tips of the wings." The ant was eaten or
discarded afterward. Pillai (1941) noticed Indian Mynahs spreading and
quivering the tail during active anting in the midst of a colony of Oecophylla
smaragdina, a spraying ant species. Tebbutt and Stone each mentioned
shaking of wings or tail in anting birds, and other instances are cited else-
where in this paper. Sick (1957) noticed that a free-living Blue-necked
Tanager (Calospiza cyanicollis) sometimes cocked its tail upward during
active anting. Some of these acts bear remarkable resemblance to certain
movements of sexually-motivated birds, as discussed by Armstrong (1947),
Hinde (1955; 1956), Moynihan (1955) and others.
Another parallel sometimes is seen between the postures of sun-bathing
and those of anting, particularly passive anting. Beebe's (MS) Northwestern
Crows sprawled on ant nests with their heads "generally held off to one side
246 THE WILSON BULLETIN September 957
Vol. 69, No. 3
to expose the fold of naked skin between the feathers of the back and
scapulars .... The nearest similar behaviour I have observed is that caused
by sudden exposure to sun when a bird has been in shade for some time."
Condry (1947) and Goodwin (1953a) described much this same posture in
anting Carrion Crows. The behavior of Burton's (1955a) Rook, already
mentioned, when exposing itself to electric heat and to steam, may have been
analogous to sun-bathing. Other sun-bathing attitudes suggestive of artting
postures can be found in Hauser (1957), Rollin (1948), and Gibb (194-7).
Hauser (op. cit.) showed that free birds, sunning themselves on a brown
masonite feeding tray or on a leafy compost heap, were exposed to surface
temperatures as high as 140 ø F. She said that heat alone did not seem to be
the primary factor. Yet her "Compulsory Sun Position," as distinct from
"Voluntary" sunning, deserves critical study in the light of birds' anting
responses to thermogenic materials. Certain of her descriptions and sketches
showing intense, involuntary sun-bathing posture seem very much like some
of the attitudes described for anting, in situations where, apparently, the
only heat involved was in the anting material itself. (Compare, for example,
Hauser's sketch of sunning Mockingbird, Mimus polyglottos, with Goodwin's
[1955b] sketch of artting Rook.)
Another point that may prove of considerable importance to better under-
standing of anting is that passive anting appears to be extremely rare, if it
occurs at all, with non-spraying ants. Except for Galloway's (1948) brief
statement that a Magpie-Lark which he saw picking up and squeezing Iri-
domyrmex detectus "did not mind the ants crawling about its feathers," all
cases I have seen of identi/ied ants invading a bird's plumage have con-
cerned spraying species. Unfortunately, Galloway did not describe the Magpie-
Lark's posture, or state whether it was applying ants or only feeding on
them. Both Goodwin (1951) and Liihmann mentioned negative response in
passively anting birds when few ants were present. This seems quite different
from actively artting birds which, in a number of instances, have been known
to respond to one or few ants; and it suggests that a considerable amount of
spray may be required for passively anting birds to assume artting posture.
It would be instructive to learn whether the passively artting Common Crow,
for instance, would respond at all to non-spraying ants.
In considering the premise that anting birds, at least in some cases, are
trying to get thermogenic materials on the ventral body skin, we should not
neglect smoke as an anting substance. IJzendoorn, Chisholm (1948:163-
175), and others, have discussed smoke in connection with antirig. It has
been suggested that smoke may be satisfying for its warmth or the thermo-
genic effects of acids contained in it. Doubtless this is true also of many
other substitute materials, such as beetles, earwigs, millipedes, wasps, pre-
pared mustard, and some of the other vegetable materials. The use of hair
Lovie M. ANTING IN BIRDS 247
Whitaker
tonic, previously mentioned, is of unusual interest, for Dr. Fluck wrote me
that the lotion contained, in addition to bay rum and alcohol, tincture of
cantharides. Cantharides are dried beetles (usually Cantharis vesicatoria,
C. vittata, or Mylabris cichorii), which have the vesicant constituent canthari-
din, as well as uric, formic and acetic acids. Of pungent, acrid taste and
penetrating, aromatic odor, these insects are used in medicine as a counter-
irritant, blistering agent, diuretic and aphrodisiac (Youngken, 1948:920;
Mansfield, 1937:463).
Some species, even individual birds, seem to perform with smoke exactly
as they do with ants and certain substitutes. I have already mentioned how
various burning or smoking materials were applied by a Rook and a European
Jay. Burton's descriptions and the photographs of these two individuals
follow closely Goodwin's descriptions and sketches of these same species
when using ants (Burton, 1955a, 1955b, 1955c; Goodwin, 1952a, 1955b).
Akhough it also used ants, Burton's Rook consistently gave strong, typical
anting responses to smoke of any kind. A captive Blue Jay, a species with
pronounced anting proclivities, applied burning cigarettes (Miller, 1952),
and another such individual held them in the bill, apparently in order to get
smoke under its wing (Anon., 1952).
A most unusual account of birds using smoke concerns the small flock
of Red-browed Waxbills (Estrilda temporalis) at Kairi, on Atherton Table-
land, North Queensland, which Givens (1945) observed on several days in
June. The birds, as many as a dozen at once, would stand on a smoldering
log, in the curling wisps of smoke coming up through cracks in the bark,
and there perform anting movements "quite distinct from those commonly
seen when birds bathe in dust or water". Each bird would stand upright,
with tail as "support" and wings drooping a little forward and downward,
and begin sweeping the head forward and down under the wings, meanwhile
vigorously shuffling its wings and body feathers and "often toppling back-
ward from the violence of its efforts." Givens, watching from a distance of
about six feet, saw no insects on the log, nor could he find any afterward.
But he does say that on one occasion "when a piece of bark was torn away
from a nearby stump," revealing an ant nest, one of the birds "tried to ant
itself there, but soon abandoned the attempt in favour of the smoke." Some-
times a bird, unable to find space in the smoke, performed a few feet away,
much as a bird may water-bathe in vacuo.
Elsdon (1948) described Linnets (Carduelis cannabina), Meadow Pipits
(Anthus pratensis) and Pied Wagtails (Motacilla alba) that for several days
in August and September persistently flew into thick smoke from a huge
oil-tank fire, at times flying as low as about 50 yards above the flames.
Sometimes birds would alight exhausted near the observers, only to fly back
into the smoke when they apparently had recovered. Jackdaws (Corvus
248 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
monedula) repeatedly have hovered briefly in chimney smoke (Ridley, 1948),
while Common Starlings and Rooks have perched in such smoke and ruffled
their feathers (Prideaux, 1947; McMeeking, 1949). One of the Rooks per-
formed "contortions" in the smoke.
These records of free birds frequenting smoke do not indicate that the
smoke was flushing out insects and thus attracting the birds, as may be the
case during grassland fires. Although some of the incidents occurred during
winter in England, others of them occurred there in June, August and Sep-
tember, when birds may not have been trying to escape cold. Among winter
records, when the insect factor would seem improbable, are some that
mention preening and posturing in the smoke.
Excepting the Linnets, Meadow Pipits, Pied Wagtails, Jackdaws and possi-
bly the Red-browed Waxbills, all birds mentioned thus far here in connection
with smoke have been species known also to use ants or the more conventional
substitutes. It is interesting to see that Poulsen (1956) found anting in the
Tree Pipit ( lnthus trivialis ) .
The other smoke-bathing records that I have seen relate to species not
known to ant: Herring Gull, Larus argentatus (White, 1948; Stevens, 1948);
Black-headed Gull, L. ridibundus (Stevens, 1948; Stafford, 1954); Little
Owl, lthene noctua (Tubbs, 1953); Swift, lpus apus (Adler, 1954); Wel-
come Swallow, Hirundo neoxena (Barker, 1939); and House Martin, Deli-
chon urbica (Pritchard, 1950).
I find no records of "smoke-bathing" as such in the Western Hemisphere;
however, Dr. Arthur A. Allen tells me that he once had a tame Common
Crow that liked to get into smoke from an incinerator, and there is the in-
stance of Bluebirds (Sialia sialis) and Cedar Waxwings (Bombycilla ced-
ro,rum), which last species is known to ant, warming themselves on a chimney-
top in sub-zero weather (Parker and Parker, 1950).
Chisholm (1948:163-175), reviewing the problems of anting, commented:
"'Smoke-bathing' may in fact be complementary to water-bathing, sun-
bathing and dust-bathing, and all four may well be allied to 'anting' with
acids."
252 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
A BIBLIOGRAPHY OF PUBLISHED REFERENCES TO ANTING
ALMA, E.
'1951 Een nfierenhoop-probleem. Natura, 48:164.
ADLER, L. P.
1951 "Smoke-bathing" of swifts. Brit. Birds, 44:281.
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1946 Audubon and anting. Auk, 63:443.
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1949 "Anting" of green woodpecker. Brit. Birds, 42:390.
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1956 Normal and irrelevant toilet behavior in Emberiza spp. Brit. Jour. Anita.
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BAGGALEY, W.
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1941a Honeyeater and ants. Emu, 41:163-164.
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1951 External parasites of birds: a review-. Wilson Bull., 63:363-369.
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254 THE WILSON BULLETIN Septenber 1957
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1948 Bronzed grackle anointing plumage with orange-skin. Wilson Bull., 60:244-
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1949 A blue jay "anting." Migrant, 20:37-38.
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'1931 Unser Korax. Oeschichte einer Rabenkrihe. Gefied. Welt, 60:425.
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1945 Chickadee notes, no. 1277. Winnipeg Free Press, Sept. 14, 1945.
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1952 Einemsen junget Eichelhiiher. Ornith. Beob., 49:28.
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1949 "Anting" of chaffinch. Brit. Birds, 42:120.
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1951 Das Einemsen bei zahm aufgezogenen jungen Kr/ihen. Jour. /. Ornith., 93:
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1914 Birds transporting food supplies. 4uk, 31:404-405.
1918 The biting power of ants. 4mer. Mus. Jo.ur., 18:141-147.
1938 "Anting" by birds. 4uk, 55:98-105.
1944 Red-eyed towhee anting. 4uk, 61:298.
1947 Wild turkey anting. 4uk, 64:130.
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1949 "Smoke-bathing" of rook. Brit. Birds, 42:22.
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1937 Materia medica, toxicology and pharmacognosy. Mosby Co., St. Louis, 707 pp.
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1948 Repeated anting by a song sparrow. 4uk, 65:600.
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'1906 The chemical nature of some insects' secretions. Bull. Wis. Nat. Hist. Soc.,
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1952 Blue jay, Cyanocitta cristata, "anting" with burning cigarettes. /tuk, 69:87-88.
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1948 La mirmecosimpatia in alcuni uccelli. Riv. Ital. Orn., 18 ser. 1I:141-144.
1952 Altri ragguagli sulla "Mirmecosimpatia" negli uccelli. Riv. Ital. Orn., 22 ser.
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1935 Relations of birds and insects. Emu, 34:248.
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1942 The nesting of African birds in association with other living things. Ibis,
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1941-1942 Observations on the nest odours of ants. Proc. Linnaean Soc. London,
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1944 A study of the ant fauna of a garden, 1934-42. Jour. Anim. Ecol., 13:123-127.
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1957 Pals of my childhood. Sat. Eve. Post, 229(41):36, 134-136, 138, (April 13,
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1955 Remarks on the original sources of displays. Auk, 72:240-246.
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1951 Red-wing, Agelaius phoeniceus, anting. Auk, 68:108.
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'1935 Werden Ameisen durch Vgel zum Vertreiben von Auszenparasiten beniitz?
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1943 Studies in the life history of the song sparrow. II Trans. Linnaean Soc. New
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1945 Cowbirds anting. Auk, 62:302-303.
1952 Breeding birds in Jackson Park. Illinois Audubon Bull., no. 82:3-7.
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1940 "Anting" by the song sparrow. Auk, 57:520-522.
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1943 "Anting" by robins. Auk, 60:5950.
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'1935 [Three young captive Cyanopica cyanus japonica applied ants on inner sides
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1950 Formic acid production among Formicidae. Ann. Ent. Soc. Amer., 43:437-443.
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1909 Strange behaviour of certain birds when in possession of strong smelling in-
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1950 Note on behavior of birds on a cold, winter day. Auk, 67:108.
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1945 Strange behavior of a bronzed grackle. Bird-Banding, 16:144.
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1954 Birds' "anting" antics. Gull, 36(11) :44.
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1938 Starlings putting ants in their plumage. Chat, 2:9.
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1930 Ectoparasites and bird-banding. Bird-Banding, 1:51-60.
1933 External parasites collected from banded birds. Bird-Banding, 4:68-75.
260 THE WILSON BULLETIN September 1957
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1936 A list of external parasites from birds of the eastern part of the United States.
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1941 Bird "bathing" in ants. Jour. Bombay Nat. Hist. Soc., 42:935-936.
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1955 Experiments on antlug by birds. Acta XI Congr. Internat. Ornith. 1954
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1956 A study of anting behaviour in birds. Dansk Ornith. Forenings Tidsskrijt,
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1951 Reactions of birds to wasps. Brit. Birds, 44:405 406.
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1947 "Smoke-bathing" of starling. Brit. Birds, 40:340.
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1950 "Smoke-bathing" by house-martins. Brit. Birds, 43:156.
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1950 Blue tits' reactions to wasps. Brit. Birds, 43:403.
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'1935 Ueber das Verhalten der Viigel gegeniiber lebenden Ameisen. Gelled. Welt,
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1948 Myrm6cop,hilie chez la Perdix bartavelle. Oiseanx, 19:288.
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1946 "Anting" by magpies. Brit. Birds, 39:313.
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1948 "Smoke-bathing" of jackdaw. Brit. Birds, 41:83.
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1945 The grackle and the green grape. Indiana Audubon Soc. Year Book, 23:14.
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1948 A note on sun-bathing in birds. Brit. Birds, 41:304-305.
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1952 Fleas, flukes and cuckoos. Philosophical Library, New York, 304 pp.
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1952 Einemsen bei einer jungen Elster. Ornith. Beob., 49:28.
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1956 The army ants. Smithsonian Report ]or 1955: 379-406.
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1946 Little shrike-thrush "artting." Emu, 46:132.
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1947 Magpies "artting." West. Australian Nat., 1(1):21.
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1947 Anting by the indigo bunting. Kentucky Warbler, 23:1.
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'1914 Beyond the pasture bars. Century, New York, 160 pp.
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1957 Anting by two tanagers in Brazil. Wilson Bull., 69(2) :187 188.
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1955 The nature of "anting." [Letter] Brit. Birds, 48:94-96.
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1948 Anting by some Costa Rican birds. Wilson Bull., 60:115-116.
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1941 "Anting" by the cardinal. Auk, 58:414-415.
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1949 Souh Carolina bird life. Univ. South Carolina Press, Columbia, 585 pp.
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1942 A robin anting. Wilson Bull., 54:214-215.
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1954 "Smoke-bathing" by black-headed gulls. Brit. Birds, 47:311 312.
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1949 "Anting" of green woodpecker. Brit. Birds, 42:59.
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'1937 "Ekelgeschmack" und visuelle Anpassung, einige Futterungsversuche an
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1948 "Smoke-bathing" of herring- and black-headed gulls. Brit. Birds, 41:244.
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1954 "Anting" by wryneck. Brit. Birds, 47:312.
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1935a Werden Ameisen durch ViSgel zum Vertreiben yon Auszenparasiten beniitzt?
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1936 Nochmals: Das Linerosen des Gefieders. Ornith. Monatsber., 44:116-117.
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1951 Birds and an ant arny in southern Tamaulipas. Condor, 53:16-18.
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1956 The perfuming act. Nature Mag., 49(7) :339.
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1953 Circle of the seasons. Dodd-Mead, New York, 306 pp.
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1946 "Anting" of starling. Brit. Birds, 39:84.
262 THE WILSON BULLETIN September 1957
Vol. 69, No. 3
TENISON, W. P. C.
1954 "Anting" by blackbird. Brit. Birds, 47:312.
THOMAS, J. W.
1957 Anting performed by scaled quail. Wilson Bull., 69:280.
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1941 "Anting" by summer tanager. Auk, 58:102.
1946 Catbird "anting" with a leaf. Wilson Bull., 58:112.
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1956 Learning and instinct in animals. Harvard Univ. Press, Cambridge, Mass.,
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'1931 Bunte Bilder aus der Vogelstube. Ge/ied. Welt, 60:484.
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1953 Little owl "smoke-bathing." Brit. Birds, 46:377.
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1943 "Anting" by the robin and towbee. Auk, 60:61.
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'1935 Ameisen und Vgel. Ge/ied. Welt, 44:419.
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1951 Line Beobachtung yon Einemsen an einer isoliert aufgezogenen Rabenkrihe
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'1943 Some experiments on the sense of smell in birds, studied by the method of
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1935 The biology of the thatching ant, Formica ru]a obscuripes Forel, in North
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1950 "Anting" of carrion crow. Brit. Birds, 43:402.
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1951 Notes on "anting." Emu, 51:81-82.
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1948 "Smoke-bathing" of starling and of herring gull. Brit. Birds, 41:244.
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1947 "Anting" by blackbird. Brit. Birds, 40:84-85.
1948 "Anting" by starlings. Brit. Birds, 41:306.
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1938 An orgy of ants. Auk, 55:679%80.
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1909 A nesting of the blue-winged warbler in Massachusetts. Auk, 26:337-345.
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1948 Phartnacognosy. Blakiston Co., Philadelphia, 1063 pp.
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1946 [in Letters.] Nat. Hist., 55:149 and 152.
1204 WEST BROOKS STREET, NORMAN, OKLAhOmA, MAY 31., 1957